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TariqHusain, BhaskarDatta 

s (Liliaceae) in Korea 

Soon Suk Rang andMyon^ 

ed Saccharum (Poaceae: Andropogoneae) 
and Kashmir, northwest Himalaya, India 
, R.R.Rao and A. Garg 


Population and site characteristics of a recently discovered disjunct populatior 
Gregory H. Aplet, Richard D. Laven, Maurya B. Falkner and Robert B. Shaw 

'. Peterson and Robert B. Shaw 

i States and Canada: 

a de la Paila, Coahuila, Mexic 

Vascular plant type specimens ii 
with a brief history of the herba 
Michael 0. Moore and David E. 

z University of Georgia Herbarium, 




Lloyd H. Shinners 


Wm. F. Mahler 
ublisher 1971-1992 
Director Emeritus 

Barney L. Lipscomb 


)tanical Research Institute of Texas, Inc. 

509 Pecan Street 

Fort Worth, Texas 76l02-4060 

817 3324441 / 817 332-4112 FAX 

John W. Thieret 

Associate Editor 

Northern Kentucky University 

Highland Heights, Kentucky 41076 

[BUTioNS TO Botany, v 

Copyrigtit 1 



The Marie Selby Botanical Gardens 

sin recognized as subger 
are presented. 

Generic level taxonomy in subtribe Gaillardiinae has long been problem- 
atic. Depending on which taxa are included in or excluded from the subtribe 
and how the included taxa are delimited into genera, subtribe Gaillardiinae 
might contain as few as two genera {Gaillardia Foug. and Helenium L.) or 
more than 25 genera. 

With regard to Dugaldia Cass, and Plummera A. Gray, questions have 
arisen as to whether they should continue to be recognized as genera or 
submerged within Hymenoxys Cass. When I resurrected Dugaldia (Bierner 
1974), those taxa were being treated as congeneric with Helenium by most 
workers (e.g., Gray 1874; Standley 1940). It has been clear for some time, 
however, not only that they are morphologically and chemically very 
different from taxa in Helenium, but that they are morphologically and 
chemically very similar to taxa in Hymenoxys (e.g., Bierner 1978). Likewise, 
the taxa included in Plummera have been recognized for quite some time as 
being very similar to taxa in Hymenoxys (Bierner 1978), and it had even been 
suggested earlier (Turner et al. 1973) that Plummera be combined with 

Consideration of these questions in light of recent micromolecular 
chemical studies (Sprmg et al. 1994) and chloroplast DNA restriction site 
analyses (Bierner and Jansen unpublished) has led me to the conclusion that 
indeed Dugaldia and Plummera should be combined with Hymenoxys and 

recognized as subgenera. For purposes of this discussion, Hymenoxys is 
considered to include taxa that at times have been segregated into the genera 
Macdougalia A. Heller, Phileozera Buckley, Pkradema Hook., and Rydbergia 
Greene, but not taxa that can be segregated as Tetraneuris Greene (see Table 1 ). 


The taxa oiDugaldia are similar morphologically to Hymenoxys brandegei 
(Porter ex A. Gray) K.L. Parker, H. grandiflora (Torr. & A. Gray) K.L. Parker, 
and H. imignis (A. Gray ex S. Watson) Cockerell, which are members of 
Hymenoxys subgenus Rydbergia (Table 1). In particular, the involucral bracts 
of these taxa are organized into two to four subequal series, and the outer 
bracts are usually densely lanate toward their bases. The leaves of the 
Dugaldia taxa tend to be entire, while those of the Hymenoxys taxa tend to be 
divided; however, Dugaldia integrifolia (Kunth) Cass, often has upper and 
middle leaves that are deeply three-toothed, and Hymenoxys brandegei some- 
times has all of its leaves entire. 

The Plummera taxa are very similar morphologically to taxa in several of the 
Hymenoxys subgenera (Table 1). These similarities include involucral bracts 
that are in two unequal series with the outer bracts united, and leaves that 
are divided into linear segments. The disc florets of Hymenoxys and Dugaldia, 
however, are hermaphroditic, while those of Plummera are functionally 


Except for reports of dysploidy in Hymenoxys odorata DC. (e.g., Sanderson 
and Strother 1973) and H. texana (J.M. Coult. & Rose) Cockerell (Strother 
and Brown 1988) of subgenus Phileozera, the taxa oi Hymenoxys , Dugaldia, 

Beaman and Turner 1962; Bierner 1974; Sanderson 1973; Speese and 
Baldwm 1952; Strother 1966; Turner et al. 1973; Table 1). No polyploidy 
has been reported in any of these taxa, but polyploidy has been reported in 
some taxa ofTi^/r^^zez^m (Johnston and Bonde 1969; Parker 1970; Speese and 
Baldwin 1952; Strother 1966, 1972). 

Flavonoid, Monoterpene, and Sesquiterpene Lactone Chemistry 

Hymenoxys, Dugaldia, and Plummera are extremely similar chemically 
(Table 1). All of them produce similar or identical 6-methoxy flavone 
aglycones, flavonol aglycones, flavonol glycosides, and 3-0-acetyl flavonol 
glycosides (Bierner 1974, 1978, 1994, unpublished; Sanderson 1975; 
Wagner etal. 1971, 1972a, 1972b). Taxa of r^/mm^m, however, have been 
found to produce 6, 8-dimethoxy flavone aglycones rather than 6-methoxy 

BIERNER, Dugaldia and Plummera in Hymenoxys 5 

flavone aglycones (Bierner unpublished; Thomas and Mabry 1967, 1968a, 
1968b), and none has been found to produce 3-0-acetyl flavonol glycosides. 

With the exception o( Hymenoxys texana (tentatively placed in subgenus 
Phileozera), monoterpene glycosides have not be detected in Hymenoxys, 
Dugaldia, or Plummera (Spring et al. 1994). Monoterpene glycosides have 
been found, however, in all of the Tetraneuris taxa (Spring et al. 1994). 

Hymenoxys (except for H. texana), Dugaldia, and Plummera all produce 
similar or identical guaianolides, pseudoguaianolides, and seco-pseudogua- 
lanolides (Bohlmann et al. 1985; Hill etal. 1977; Ivieet al. 1976;Romode 
Vivar et al. 1987; Seaman 1982; Spring et al. 1994). Tetraneuris taxa again 
are somewhat different; they produce guaianolides and pseudoguaianolides, 
but seco-pseudoguaianolides have not yet been found (Seaman 1982; Spring 
etal. 1994). 

Chloroplast DNA 

Recently, I worked with Robert K. Jansen at the University of Texas at 
Austin on chloroplast DNA restriction site analyses of many of these taxa 
(this work is being prepared for separate publication). Subgenera Hymenoxys 
and Rydbergia were not represented, but one species o( Dugaldia (D. hoopesii 
[A. Gray} Rydb.) and both species of Plummera were included. In the 
phylogenetic analysis of the data, the Dugaldia and Plummera taxa were very 
strongly supported as being in the same branch of the chloroplast DNA 
phylogenetic tree with the taxa of Hymenoxys subgenera Macdougalia, 
Phileozera, and Picradenia, while the taxa o( Tetraneuris were separated with 
strong confidence into a separate branch. 

Hymenoxys Cass., Diet. Sci. Nat. 55:278. 1828. Type Species: Hymenopappus 
anlhemoides Juss., Ann. Mus. Natl. Hist. Nat. [Pans] 2:426. 1803. 

Hymenoxys subgenus Dugaldia (Cass.) Bierner, stat. nov. Basionym: Dugaldia 
Cass., Diet. Sci. Nat. 55:270. 1828. Type Species: Aainea integrifolia Kunth. 
Oxylepts Benth., PI. Hartw. 87. 1841. Type Species: Oxylepis lanata Benth. 

Hymenoxys integrifolia (Kunth) Bierner, comb. nov. Basionym: Actmea 
integrifolia Kunth, Nov. Gen. et Sp. 4:297. t. 410. 1820. Dugaldia integrifolia (Kunth) 
Cass., Diet. Sci. Nat. 55:270. 1828. Cephalophora integrifolia (Kunth) Steud., Norn, 
ed. 2. 1(3):328. 1840. Helemum integrifolia [sic] (Kunth) Benth. & Hook, ex Hemsl., 
Biol. Centr. Amer. Bot. 2:227. 1881. Heleniastrum integrifolium (Kunth) Kuntze, 
Revis. Gen. Pi. 1 :342. 1891 . Type: MEXICO. Hidalgo: "...inter Omitlan et Serro del 
Jacal, alt. 1400 hex." Humboldt & Bonpland s.n. (holotype: P!). 
Oxylepis lanata Benth., PI. Hartw. 87. 1841. Helenium lanatum (Benth.) A. Gray, Proc. 

Amer. Acad. Arts 9:205. 1874. Type: GUATEMALA: "In summo monte Cumbre de 

Argueta dicto," Uartweg 593 (holotype: K!; isotype: P!). 

Hymenoxys hoopesii (A. Gray) Bierner, comb. nov. Basionym: Helemumhoopesn 
A. Gray, Proc. Acad. Nat. Sci. Philadelphia 1863:65. 1864. Helemastru?n hoopesii (A. 
Gray) Kuntze,Revis. Gen. PI. 1:342. 1891. Dugalciia hoopesn (A.Gmy)Rydh., Mem. 
New York Bot.Gard. 1:425. 1900. Type: UNITED STATES. Colorado: "South Park 
and west of Pike's Peak," Ha// & Harbour 272 (holotype: GH!; isotypes: MO!, NY!, 

Hymenoxys pinetorum (Standi.) Bierner, coml 

pinetorim Standi., Field Mus. Pub. Bot. 22:127. 1940 
Bierner, Bnttonia 26:391. 1974. Type: MEXICO. 
Oriental, ascent of Sierra Potosi by north hogback. 

The species of Hymenoxys subgenus Dugaldia are 
already been treated taxonomically (Bierner 1974). 

Hymenoxys subgenus Plummera (A. Gray) Bierner, stat. nov. Basionym: 
F/ummera A. Gray, Proc. Amer. Acad. Arts 17:215. 1882. Type Species: P/ummera 
fiori/junda A. Gray. 

Hymenoxys microcephala Bierner, nom. nov, Basionym: P/ummerafioribundaA. 
Gray, Proc. Amer. Acad. Arrs 17:215. 1882. non Hymenoxys floribunda (A. Gray) 
Cockerell, Bull. Torrey Bot. Club 31:485. 1904. Type: UNITED STATES. Arizona: 
Cochise Co.: "Apache Pass, Chirricahua [sic] Mountains," Sep 1881, Lemmon & 
Lemmon 332 (holotype: GH!; isotypes: BM!, F-207616!, GH!, K [photo at F- 
1645644!], NDG-061757!, PH-two sheets!, US-47542!; Probable isotypes: F- 
313722!, MO-3726424!, NDG-06I758!, NY!). 

22 Jul 1927, Peeb/es, Harrison & Kearney 4393 (holotype: US-1436073!, photo at 
The two taxa oi Hymenoxys subgenus Plummera could be treated as varieties 
of a single species, as suggested by Turner et al. (1973). I agree that 
morphologic differences between the two are slight, but the taxa appear to 
be geographically isolated from one another without a zone of contact and 
intergradation. Furthermore, Spring et al. (1994) found several differences 
between them with regard to sesquiterpene lactone substituents and side 
ive approach of continuing to recognize them 
I appropriate. 

I thank the curators of the various herbaria from which type specimens 
were borrowed. This work was supported in part by a National Science 
Foundation Research Opportunity Award supplement to grant BSR-9200707 
(Robert K. Jansen). 

.EAMAN, J.H. and B.L. Turner. 1962. Chromosome numbers u 

Compositae. Rhodora 64:271-276. 
iiERNER, M. W. 1 974. A systematic study oiDugaldia (Composit 
. 1978. Flavonoids oiFlummera: taxonomic impii 

within the Gaillardimae. Biochem. Syst. Ecol. 6:293-295. 
1994. Pectoiinarigenin from Dugaldia pinetorum ( 

Syst. and Ecol. 22:109-110. 
BoHLMANN, R, L.N. iVTiSRA and J. Jakupovic. 1985. Seco-helenanolides from 
integnfolta. Phytochemistry 24:1378-1380. 

Amer. Acad. Arts 9:187-218. 
Hill, D.W., H.L. Kim, C.L. Martin and B.J. Camp. 1977. J. Agric. Food Chem. 

I VIE, G.W., D.A. WiTZEL, W. 

Hymenovin from Hymenoxys 

weed). J. Agric. Food Chem. 24:681-682. 
Johnston, M.A. and E.K. Bonde. 1969. In lOPB chromosome reports XXII. Taxon 

Parker, K.R 1970. Two new taxa in Texas Hymenoxys. Phytologia 20:192. 
RoMO de Vivar, a., G. Delgado, M. Soriano-Garcia, R.A. Toscano, A. Ruben, E. Huerta 

and R.G. Reza-Garduno. 1987. Sesquiterpene lactones from a population oi Helenium 

tntegnfoltum.}. Nat. Prod. 50:284-286. 
Sanderson, S.C. 1973. In lOPB chromosome reports XL. Taxon 22:285-291. 

\ L.E. Brown. 1988. Dysploidy in Hymenoxys texana (Coi 

J. Bot. 75:1097-1098. 
THOMAS,M.B.andT.J.MABRY. 1967.Isolati 
flavone from Hymenoxys scaposa. J. Org. < 

Turner, B.L., A.M. Powell and T.J. Watson. 197.3. Chromosome nur 

Asteraceae. Amer. J. Bot. 60:592-596. 
Wagner, H., M.A. Iyengar, E. Michahelles and W. Herz. 1971. Querc 

(3-D-glucopyranoside in Plummera ftoribunda and Helenium hoopesii 

iiscry 11:3087-3088. 



P.O. Box 671327 

Chugtak, AK 99367-1327, U.S.A. 


Department of Biological Sciences 

California State University Chico 

Chtco, CA 95929, U.S.A. 

urn Columbian. 

urn Nutt. IS a large and morpholo, 



ible complex of popul; 

ending from 

southern Canada to northern M 

exico, i 

ind 1 

ilbiferous pc 

.pulatjons have a restricted distr 


1, oc 

currmg in the Cascac 

)m northern 1 

to southern Oregon, m the Kiamat 



IS of northern Californi 


anct group in the CahformaS:erra 


a sot 


>us aconites < 

ire probably the most distinctive 

? group 


hi n the A. columbianu 

and they an 

, treated herein as a subspecies. 

I Nutt. es un complejo grande y variable morfologicamente c 
enden desde el sur de Canada hasta el norte de Mexico, y desc 
'ork. Laspoblaciones bulbiferas tienen una distribucion restringid; 
lillera de la Cascada desde el norte al sur de Oregon, en las Montaiii; 
ilifornia, y un pequefio grupo disyunto en al sur del Lago Tahoe e 

Aconitum columbianum Nutt. is a polymorphic complex of populations 
occurring in the mountains of western North America from British Colum- 
bia, Canada to Chihuahua, Mexico and from California to South Dakota. 
Disjunct, outlying populations of A. columbianum occur also in the Driftless 
area of Iowa and Wisconsin (Brink 1 982), and in Ohio and New York (Brink 
& Woods submitted for publication; litis 1965). Previous investigations 
indicated extreme differences among regional groups of populations, par- 
ticularly in floral morphology and other correlated characters. This variation 
was continuous, however, with no clearly defined breaks, if populations 
throughout the range of the species were considered in aggregate (Brink 

1975, 1980, 1981, 1982; Brink & deWet 1980). Bulbiferous populations 
were somewhat different in this regard since they appeared to be consistently 
distinct from all other A. columbianum. The purpose of this investigation was 
to determine the geographical distribution of these bulbiferous aconites, 
and to review the taxonomy of this group. 


Aconitum populations of the United States were compared in field, 
herbarium, laboratory, and uniform nursery investigations. Geographical 
distribution of bulbiferous populations was determined by field investiga- 
tion and by mapping locations from herbarium specimens (CAS, CHSC, 
CLM, ILL, JEPS, OSC, RM, UC, US, UTC, WILLU, WS). Acomtum plants 
from bulbiferous and non-bulbiferous populations were transplanted and 
grown in growth chambers and greenhouses to study bulbil production 
under controlled conditions. 

In field investigations we found no intergradation between bulbiferous 
and non-bulbiferous Aconitum columbianum (however, see Hitchcock et al. 
1964; Abrams 1944). The populations that we have seen were either 
bulbiferous, with all plants of sufficient size and maturity producing 
conspicuous bulbils in the leaf axils, or they were completely non-bulbifer- 
ous with no bulbil producing plants. In growth chambers and greenhouses, 
only plants from bulbiferous populations produced bulbils. Bulbils were 
illustrated in Brink (1980) and Hickman (199.3). 

Bulbils are an effective means of vegetative reproduction. In natural 
populations they fall to the ground late in the season where they sprout 
vigorously, giving rise to new plants. A single leaf axil may produce one or 
more bulbils. Bulbil production usually is greatest at about the middle of 
the aerial stem, and diminishes toward the top and bottom of the plant. Most 
bulbiferous plants bear both bulbils and flowers. On a small percentage of 
plants, however, bulbifery completely supplants flower production, and 
bulbils are produced to near the apex of the terminal inflorescence, in place 
of flowers. A more in-depth description of bulbifery can be found in Brink 
(1975). Bulbil production should not be confused with the production of 
one to several small daughter tubers at the first few nodes above the parent 
tuber, usually below ground. This occurs on a small percentage of the plants 
in all bulbiferous and non-bulbiferous populations. 

Bulbiferous Acomtum columbianum have a restricted distribution (Fig. 1), 
with one group of populations occurring in the Sierra Nevada mountains of 
California south of Lake Tahoe in El Dorado, Amador and Alpine counties; 

and another group beginning ca. 350 miles away in northern California in 
the Klamath Mountains of Siskiyou and Del Norte counties. Bulbiferous 
populations extend from southern to northern Oregon in the Cascade range 
in Josephine, Jackson, Klamath, Douglas, Lane, Deschutes, Linn, Jefferson, 
Marion, Clackamas, Wasco, and Hood River counties. 

Munz and Keck (1968) were uncertain whether the two disjunct groups 
of bulbiferous Acomtum in California (Fig. 1) should be treated as separate 
taxa. We can discern no consistent morphological features separating these 
two groups (Brink 1975). 

We have decided to treat bulbiferous Acomtum columbianum at the 
subspecific level, due in part to the nature of the variation within the species 
as a whole. The A. columbianum complex is extremely polymorphic (Brink 
1980, 1982; Brink & de Wet 1980). The existence of geographical races led 
to a proliferation of published names at the specific, subspecific and varietal 
levels. This nomenclatural proliferation was often the result of regional 
studies, which did not consider patterns of variation within the entire 
Acomtum columbianum complex. Geographical races that appear quite dis- 
tinct within a regional context may be indistinguishable from other races 
occurring hundreds or even thousands of miles away. Robust and diminutive 
races that are regionally distinct are "connected" by intermediate races if the 
entire range of the species is considered (Brink 1980, 1981, 1982). 

We have treated Acomtum columbianum as one, large, polymorphic, 
intergrading complex. Only one group of populations is consistently 
distinct from all others. These are the bulbiferous aconites, which occur in 
California and Oregon. We have chosen to distinguish this group as a 
subspecies because it is geographically and morphologically distinct. Rec- 
ognition of these bulbiferous aconites at the species level would not seem to 
be warranted, especially since we have not accorded formal taxonomic 
recognition to regional variants that are exceedingly diverse, but in charac- 
ters that intergrade throughout the range of the complex (Brink 1982). 

Aconitum columbianum Nutt. in Torrey and Gray, Fl. N. Amer. 1:34. 
Tuber to ca. 6 x 1.5 cm; "parent" tuber produces 1 (rarely 2) "daughter" 
tubers with a connective so short the tubers are essentially contiguous. Stem 
erect and stout to twining and reclining, 2-30 dm long. Leaves 5-15 cm 
broad, deeply 3-5 (7) cleft, lobes variously incised and toothed. Inflores- 
cence racemic, terminal, or terminal and axillary. Flowers typically deep 

, Woods a 

purple to blue, occasionally white, yellowish, or blue-tinged; 18-50 i 
high, helmet 11-34 mm high, pendent sepals 6-16 mm long. 

la. Aconitum columbianum subsp. columbianum 

Aconitum arizonkum Greene; A. bakeri Greene; A. cheirophyllum Greene; A. columbianum 

ochroleucumA. Nelson; A . columbianum suh?,p. pallidum Piper; A . divaricatum Rydberg; 
A. geranioides Greene; A. glaberrimum Rydberg; A. gracilentum Greene; A. helleri 
Greene; A. infectum Greene; A. insigne Greene; A. leibergii Greene; A. lutescens A. 

noveboracensevzv. quasiciliatum Fassett; A. ochroleucum Rydberg; A. obtusiflomm Greene; 

A. patens Rydberg; A. porrectum Rydberg; A. ramosum A. Nelson; A. robertianum 

Greene; A. sukaesium Greene; A. tenue Rydberg; A. tricorne Greene; A. uncinatum 

subsp. noveboracense (A. Gray) Hardin; A. vestitum Greene. 
Spring-fed bogs, seep areas, meadows, along streams, and in other wet 
areas in the mountains of western North America from British Columbia, 
Canada to Chihuahua, Mexico at elevations of ca. 900-4,000 m. Brit. Col., 
Wash., Ore., Calif, Nev., Mont., Idaho, Utah, Ariz., S. Dak., Wyo., Colo., 
and N. Mex. Disjunct, outlying, populations occur at lower elevations (as 
low as 300 m), in Iowa, Wisconsin, Ohio and New York. 

Chromosome Number: n = '^ [Wiens & Halleck 1962, Cates 1968}, 2n = 
16 [Longacre 1942, Kawano 1965, Crawford & Gardner 1974, Brink 1975], 
2;z = 18 [Crawford & Gardner 1974}. 

lb. Aconitum columbianum subsp. viviparum (Greene) Brink comb, 
nov. Basionym: A. viviparum Greene, Feddes Repert, Spec. Nov. Regni Veg. 7:2. 
1909. Type: OREGON: swamps near Crater Lake, southern Oregon, Cmick 2912 
(Lectotype, designated herein: WS!; isolectotype: UC!). 
Aconitum bulhtferumWo^fiW, Fl. NW Amer. 1:25. 1897. Non A. Wi-/>ra»2Reichenbach, 
Ubers. Aeon. 55. 1819. A. howellii Nels. & Macbr., Bot. Gaz. 56:473. 1913. A. 
columbianum var. howellii (Nelson & Macbride) C.L. Hitchcock, Vase. Pis. Pacif. NW 
2:231. 1964. OREGON: marshes eastern slope Cascade Mountains near Mt. Hood, 
A. hanseni Greene, Feddes Repert. Spec. Nov. Regni Veg. 7:3. 1909. CALIFORNIA. 
Amador Co.: Hope Valley, 1892, George Hansen. [Note: Hope Valley is actually in 
Alpine County.] 
A. platysepalum Greene, Feddes Repert. Spec. Nov. Regni Veg. 7:2. 1909. CALIFOR- 
NIA: S side Mt. Shasta, northern California, Jul 1897, H.E. Brown. 
Spring-fed bogs, seep areas, meadows, along streams, and in other wet 
areas at elevations of ca. 900-2,500 m in the Cascade Range from norrhern 
to southern Oregon, in the Klamath Mountains of northern California, and 

with a small disjunct group of populations in the California Sierra Nevada 
south of Lake Tahoe. 

These bulbiferous aconites were first distinguished as Acomtum hulbtferum 
Howell in 1 897 , but this name is rejected as a later homonym of A . bulbtferum 
Reichenbach, which is part of A. vartegatum L., a European aconite. Three 
specific names were published simultaneously in 1909 by E.L. Greene for 
bulbiferous Acomtum. One of these names (A . vtviparum), was chosen to serve 
as the basionym for the new subspecific combination, which refers to all 
bulbiferous Aconitum columbianum. The lectotype was chosen from among 
the syntypes of the basionym. 

Bulbiferous populations in California have small flowers and shallow 
nectaries (Brink 1975), and we suspected that this would be the case for all 
bulbiferous Aconitum columbianum. We found, however, that bulbiferous 
aconites in the Mount Hood region of Oregon, at the northern extreme of 
the range of bulbiferous populations, have relatively large flowers and deep 

Flower color in A. columbianum is typically deep purple to blue; however, 
color variants include completely white, yellowish, cream-colored, or blue- 
tinged at the edges of the sepals. Color variants occur occasionally within 
blue-flowered populations, but there are also white-flowered populations, 
and groups of populations. For example, a group of white-flowered, non- 
bulbiferous populations occurs in northeast Oregon and southeast Washing- 
ton. Consequently, it is not surprising that there are also white-flowered 
bulbiferous populations, which occur in Alpine and Amador counties of 
California (see Representative Specimens, below). 

nber: 2n = 18, 19, 20 [Brink 1975}. 

Alpine Co.: Hope Valley, 29 Aug 1974, Brink & Mayer s.n. (CHSC) [flowers white, blue- 
tinged at edges]. Amador Co.: E of Silver Lake Dam, l4 Aug 1982, McNeal 2730 (OSC) 
{white-flowered]. Del Norte Co.: Dunn Creek, 6 Aug 1938, Keck 4793 (DS). El Dorado 
Co.: Lily Lake, 12-14 Aug 1978, Brink 1469 (CHSC); Echo Summit, 29 Aug 1974, Brink 
& Mayer s.n. (CHSC). Siskiyou Co.: Red Rock Creek, 26 Jul 1974, Brink & Brink s.n. 
(CHSC); Kangaroo Lake and vicinity, 24 Jul 1974, Brink s.n. (CHSC). OREGON. 
Clackamas Co.: Clackamas Lake, 24 Jul 1927, Peck 15852 (DS). Deschutes Co.: Tumalo 
Creek, 12 Aug 1906, Whued 3162 (OSC). Douglas Co.: Diamond Lake, 17 Jul 1924, 
Applegate4l34 (OSC). Hood River Co.: Barlow Pass, Mount Hood, 3 1 Jul 1934, Thompson 
11208 (US). Jackson Co.: WoodruffMeadows, 26jul 192^, Pendleton s.n. (OSC). Jefferson 
Co.: Camp Sherman, 27 Aug 1962, Schoth 1929 (OSC). Josephine Co.: Lake Creek, 
Siskiyou Mountains, 18 Jul 1949, Whittaker SS217 (WS). Klamath Co.: Pole Bridge, 
Crater Lake Park, 31 Jul 1922, Applegate 3413 (UC). Lane Co.: Gold Lake, 1 Sep 1962, 
Dennis 2433 (UTC). Linn Co.: Monument Peak, 9 Aug 1947 , Alters. n. (OSC). Marion Co.: 
Olallie Meadow, 9 Aug 1978, Dawn 63 (OSC). Wasco Co.: Barlow Road, 1 Sep 1 964, French 
2604 (OSC). 

We gratefully acknowledge anonymous reviewers, J. Bissell, L. M, Mayer, 
J.M.J. deWet, R.A. Schlising, the herbaria from which loans were obtained 
(JEPS, OSC, UC, WILLU, WS), and the herbaria that were visited. We 
appreciate receiving loans through the herbarium of the Cleveland Museum 
of Natural History (CLM). 

Univ., Chico. 

1980. Reproduction and variation in Aconitum (Ranuncuiaceae), wit! 

emphasis on California populations. Amer. J. Bot. 67:263-273. 

(Ranuncuiaceae) of the United States. Ph.D. diss. Univ. of Illinois, Urbana. 
1982. Tuberous Acomtum (Ranuncuiaceae) of the continental United States 

morphological variation, taxonomy and disjunction. Bull. Torrey Bot. Club 109:1 3-23 
and J.M.J. De Wet. 1980. Interpopulation variation in nectar production ir 

and J.A. Woods. Acomtum (Ranuncuiaceae). Flora of North America 

(submitted for publication), 

Intermountain Region. Master's thesis. Utah State Univ., Logan. 
Crawford, D.J. and R.C. Gardner 1974. Documented plant chromosome numben 

1974:2. Sida 5:292-294. 
Hickman, J.C. (Ed.) 1993. Thejepson manual: Higher plants of California. University ol 

California Press, Berkeley. 
Hitchcock, C.L.A., A. Cronquist, M. OwNBEvandJ.W. Thompson. 1964. Vascular plants 

of the Pacific Northwest. Vol. 2. Univ. of Washington Press, Seattle. 
Iltis, H.H. 1965. The genus Gentianopsis (Gentianaceae): Transfers and phytogeographic 

comments. Sida 2:129-154. 
Kawano, S. 1965. Application of pectinase and cellulase in an orcein squash method. Bot. 

Mag. (Tokyo) 78:36-42. 
LoNGACRE, D.J. 1942. Somatic chromosomes oi Aconitum noveboracense and A. uncinatum. 

Bull. Torrey Bot. Club 69:235-239. 
MuNZ, PA. and D.D. Keck. 1968. A California flora and supplement. Univ. of California 

Press, Berkeley. 
WiENS, D. and D.K. Halleck. 1962. Chromosome numbers in Rocky Mountain plants. 1, 

Bot. Not. 115:455-464. 


DSCHER, Kelly. 1992. Medicinal Wild Plants of the Prairie. An 
Ethnobotanical Guide. (ISBN 0-7006-0527-4, pbk.) University 
Press of Kansas, 2501 West 15th Street, Lawrence, KS 66049-3904. 
$25.00 (hbk), $9.95 (pbk). 340 pp. 

;s, Indian names, scientific name, description, habitat, parts used, Indian use, Anglo 
ise, medical history, scientific research, and cultivation (not all of these for every plant). 
1-page line drawing illustrates each of these species. The second part, not illustrated, 

dex. Data in the book are mainly historical; only a few of the species have been studied 
itly to learn what biologically active compounds they contain.— ^o/?« W. Thieret. 

sowsK], SALLYand Andy Wasowskl 1992. Requiem for a Lawnmower. 
(ISBN 0-87833-81 1-X, hbk.) Taylor Publishing Company, 1550 
West Mockingbird Lane, Dallas, TX 75235. $15.95 (hbk). 182 pp. 
lis book, written "with Andy Wasowski," is a series of essays about gardening with 
'e plants. The book has a bias toward the Southwest, but the principles presented can 

gardeners. — -John W. Thieret. 

KiRKPATRiCK, ZoE Merriman. 1992. Wildflowers of the Western Plains. 
(ISBN 0-292-79061-9, hbk.) University of Texas Press, Box 7819, 
Austin, TX 78713-7819. $24.95. 240 pp. 

Tveten,John and Gloria TvETEN. 1993. Wildflowers of Houston. (ISBN 
0-89263-319-0, pbk.) Rice University Press, PO. Box 1892, Hous- 
ton, TX 77251. $18.50 (pbk), $29.85 (hbk). 309 pp. 

plant, flower, and fruit; notes on range; and remarks. The composites occupy nearly 1/4 of 
the book, attesting to the prevalance of membets of that family among at least the more 
showy flowers of the region. Wildflowers of Houston includes about 200 species grouped by 
flower color, briefer descriptions than Western Plains, and remarks. Both volumes are 
splendidly illustrated with color photographs of high quality. Both have a bibliography and 



Taxonomy & Herbarium Division 

National Botanical Research Institute 

Lucknow-2 26001, INDIA 

griffithtana var. pallida. 

While examining type specimens of Berberis on loan from the British 
Museum (BM), the authors came across two sheets from Tibet, both 
annotated by CM. Hu as Berberts grtffithiana Schneid. var. pallida (Hk.f. & 
Th.) Chamberlain & Hu and were included in the systematic treatment of 
Berberis by Chamberlain & Hu (1985). Critical examination of these two 
type sheets revealed considerable differences between them. One sheet 
{Ludlow & Sherriffl274) was annotated by Ahrendt as 6. replicata W.W. Sm. 
var. dispar Ahrendt (Fig. 1), and the other sheet {Ludlow, Sherriff 8c Elliot 
12518) annotated by Ahrendt as B. atrocarpa Schneid. var. trimensis Ahrendt 
(Fig. 2). 

Chamberlain c& Hu (1985) separated the two species, Berberis replicata and 
B. griffithiana as below: 

Further examination of L. & S. 1274 revealed all characters oi Berberis 
replicata whereas L. S. 8c E. 12318 clearly revealed the characters of B. 
griffithiana. Even Ahrendt (1961) had kept B. replicata var. dispar under B. 
griffithiana without citing this particular type specimen under this taxon. 

Further, the study of floral characters reveals that the apices of the stamens 
are truncate with long connectives in L.S.&E. 12318 while such character 
IS not noticed in L. & S. 1274. The gland on the petals of L. S. & E. 12318 
are ovoid, 1.25 mm long, with rounded apices, whereas inL. 8c S. 1274 the 
glands on the petals are distinctly ovate, much longer, and with acute apices. 

■t of Berheris rep/k. 

r {Ludl 

These characters clearly support the separation of these two t 
The evidence suggests that Chamberlain & Hu (1985) v 
reducing B . taromnm var. trimensh to a synonym of B . griffithk 


HusAiN, Datt and Rao, Bej 

FIG. 2. Berbens griffithiana var. pallida {Ludlow, Sherriff & Elliot iH 125 

Further, data through SEM studies of the epidermis of mature leaves of the 
wo taxa also revealed significant differences (Fig. 3), supporting the above 
'iew. In case of Berberis replicata var. dispar (Fig. 3, A) the lower epidermal 
ells of leaves exhibited the convex nature of the periclinal wall cells, with 
note or less regular cell boundaries and smooth cuticle, whereas in case of 





FIG. 3 Scanning electron micrographs of the abaxial surface of the leaf, 


replicata var. dispar (Ludlow & Sherriff 1274, BMj; (B) Berberis griffithiana 


(Ludlow. Sherriff & Elliot 12318, BMj, Bar = 10 pm. 

B. griffithiana m^'l. pallida (Fig. 3, B) the periclinal wall cells ar 

e concave 

nature, with irregular cell boundaries and the cuticle is more oi 

r less striat 

The number of stomata also varied in the two species. The forr 

nerwith t( 

stomata m one field and the latter with six. 

Berberis repltcata var. repltcata, B. replkata var. dispar and B. griffithiana \ 
allida are separated by the morphological characters in Table 1 . 


The authors are thankful to the Director, National Botanical Research 
Institute, Lucknow, for facilities and to the British Museum of Natural 
History (BM), and the Royal Botanic Garden, Edinburgh (E), for loan of 

, D.F. and CM. ] 


ARD, RobertI. 1993. Guide to the Grasses ofthe Lower Rio Grande 
Valley, Texas. (ISBN 0-938738-08-9, pbk.) University of Texas-Pan 
American Press, Edinburg, TX 78539- Price not given. 240 pp. 

Barbour, Michael, Bruce Pavlik, Frank Drysdale and Susan Lindstrom. 
1993. California's Changing Landscapes: Diversity and Conser- 
vation of California Vegetation. (ISBN 0-943460-17-4, pbk). Cali- 
fornia Native Plant Society, Sacramento, CA 95814. $24.95. 224 pp. 
Gary Snyder, the wise C^alifornia poet, in the preface notes the "elegance and economy" 

with which this important book makes California's "vast and complex landscape available 

do not become entirely degraded, and whose grasslands and forests are sustainable. " It is very 
significant that the authors address the landscape of vegetation and the natural dynamics 
that both sustain those landscapes and are perturbed by human influences. Because it is from 
that perspective, starting with understanding, that we can develop the management 
practices to protect, restore, and maintain native biodiversity. In this way the book is a model 
to lead other states, a role taken by the California Native Plant Society, publishers of this 
book, and several partner conservation organizations for several decades. 

The book is readable and informative, for both lay and technical audiences. An 
introductory chapter describes the environmental forces that have shaped California's 
landscapes as well as the changes that have occurred since European settlement. The 

and diagrams. Two closing chapters raise new and important topics: the first on past 
influences on the vegetation by Native Californians, previously thought to have lived for 
many millennia in California's lush vegetation without significant influence on its structure 
or function; the second on attempts and opportunities to restore degraded places, a goal that 

■aged by Gary Snyder. — Larry For, 



Department of Biology 

Gyeongsang National University 

Chinju 660-701, THE REPUBLIC OF KOREA 

sis shows that the type of Hemerocallis micrantha Nakai is 
e of morphological variation of populations off/, hakuunemh 
Drthwestern Korea). Hemerocaltis hakuunensis has priority and 
an. Also, the result shows that H. middendorffii (central and 

: perianth tube and a different flowering period. Synonymy, 

mponentes principals muestra que el tipo de Hemerocal 
range continuo de variacion morfologica de las poblacior 
y noroeste de Korea). Hemerocallis hakuunensis tiene prior 

Hemerocallis L. is an economically important genus of approximately 30 
species restricted to mainly eastern Asia (Matsuoka and Hotta 1 966). Many 
species and cultivars are widely grown in gardens in Asia, Europe, and North 
America (Cohen 1 986). Numerous nomenclatural and taxonomic problems 
exist within the genus (Matsuoka and Hotta 1966; J. Noguchi pers. comm.). 
The taxonomic difficulty has been attributed to the relative paucity of 
diagnostic characters, the fact that many species (e.g., H. aurantiaca Baker, 
H. flava L., H. fulva L., and H. thunbergii Baker) were described from 
cultivated plants of unknown origin (Kitamura et al. 1986; Matsuoka and 
Hotta 1966), the extreme difference in appearance between living plants 
and dried herbarium specimens, and possible widespread hybridization 
(Kitamura et al. 1986). In addition, many species o^ Hemerocallis are so 
variable ecologically and morphologically that a proper species concept 
requires morphological, ecological and biosystematic studies. Over 27,000 
cultivars further confound the taxonomic status of several Hemerocallis 

species. Matsuokaand Hocta(1966) noted that Baily (1930), Nakai (1932), 
and Stout (1941) did not consider the variability of natural populations when 
developing their classification. Based on the taxonomic literature on the 
genus Hemerocaiiis {e.g., Naka.i 1932; Matsuokaand Hotta 1966; Hotta 1986; 
Kitamura et al. 1986), the geographical and ecological distribution of the 
Japanese Hemerocallis is relatively well known. On the other hand, little is 
known of the Korean Hemerocallis. After Nakai's (1932) description of H. 
coreana Nakai as the endemic Hemerocallis species in Korea, he (Nakai 1943) 
described two other taxa; H. hakuunensis Nakai and H. micrantha Nakai. 
Matsuoka and Hotta (1966) recognized H.fulva var. minor (Miller) M. Hotta, 
H.flava var. coreana (Nakai) M. Hotta, H. dumortieri Morren var. middendorffii 
(Tr. et Mey.) Kitamura, and H. hakuunensis as the indigenous species in 
Korea. Lee (1985) reported 6 species and two cultivated plants from Korea: 
H.fulva L. (cultivated plant), H.fulva var. kawanso Kegel (cultivated plant), 
H.flava, H. littorea Makino, H. dumortieri, H. middendorffii, H. minor znd H. 
thunhergii. The recognition of Korean Hemerocallis species has varied depend- 
ing on authors. Consequently, there is an obvious need to study the group 
in detail to understand better the taxonomy of Hemerocallis. Principal 
components analysis of morphological data has been widely used to study 
species delimitation; e.g., Noguchi (1986) for the Hemerocallis dumortieri 
complex and Chung et al. (1991) for the Korean Hosta (Liliaceae). 

The goals of this study were to 1) analyze morphological variation 
between H. hakuunensis and H. middendorffii; 2) determine the overall 
geographic distribution patterns for H. hakuunensis and H. middendorffii; and 
3) more closely conform the descriptions to the typical morphology of H. 
hakuunensis. More specifically, do the two species H. hakuunensis and H. 
micrantha merit taxonomic distinction at the rank of species? 


Several morphological characters were measured for five individuals from 
16 populations in Korea from 1988 to 1993 (Fig. 1; Table 1). Observations 
were also made of the habitats, and notes were taken on characteristics (e.g., 
soil types) of the populations. Voucher specimens of all collections are 
deposited at GNUC. In addition, five herbarium specimens of H. middendor- 
ffii and the two types of H. hakuunensis and H. micrantha (Table 1) were used 
to determine if 1 ) the types of the latter two names fall within one continuous 
field of variation and 2) H. hakuunensis and H. micrantha are morphologically 
distinct by comparison with H. middendorffii. The herbaria of Seoul National 
University (SNU) and Tokyo University (TI) were visited in order to search 
type specimens off/, hakuunensis and H. micrantha and determine the total 
range of geographic variation and the distribution of North Korean Hemero- 
callis (Fig. 1). 

1. Geographical distribution oi Hemerocallis hakuunensis and H. middendorffii 
Arrows with alphabetic codes (A-N, P and Q) indicate locations of populations fro 
samples were collected by M. G. Chung and S. S. Kang. Arrows with codes (R-] 

Ltionof23 alphabetic codes. 

E Pyeongmok-ri, Imwhe-myeon, Chindo-gun, Chin Island, Chollan 

F ClVosang-n, Uishin-myeon, Chindo-gun, Chin Island, Chollanam 

H Yongdam-n, Kumnam-m'yeon, Yeongi-gun, Ch'ungch'ongnam-d( 

I Changg.-ri, Anmyeon-eup, Seosan-gun, Chungch'ongnam-do 

L Yuljeon-ri, Die-myeon, Hongch'on-gun, Kangwon-do 

N Sangbuk-myeon, Uljoo-gun, Gagisan Provincial Park, Gyrongsan 

'ungjin-shi, Hamgyeongbuk-do 

:. Kwanmyobong, Gyeongsung-gun, Hai 

Previous studies (Matsuolca and Hotta 1966; Hotta et al. 1966; Hotta 
1986; and Noguchi 1986) revealed that several qualitative (e.g., flowering 
time and period, shape of roots, odor, and type of an inflorescence) and quanti- 
tative characters (e.g., floral and leaf morphologies and wintering conditions 
of leaves) are important characters in recognizing Hemerocallis taxa. As most 
qualitative characters were not available from the herbarium specimens, 
only 14 morphological characters were selected in this study (Table 2). 

Phenetic analyses were utilized to assess the morphological variation 
present in 87 OTUs (operational taxonomic units) of H. hakuunensis, H. 
mkrantha, and H. mtddendorffii . Principal components analyses (PCAs) were 
performed to identify characters contributing to the separation of each taxon 
and assess inter-taxon variation using the Statistical Analysis System (SAS 
1987). The data were standardized for each character, with the raw data 
matrix transformed such that each character had a mean of zero as a standard 
deviation (Sneath and Sokal 1973). The first three principal components 
were extracted by the PCAs. 

The cumulative variance of the first three principal components (64.8%) 
and the contributing value of each of 14 characters are shown in Table 3- 
Characters weighted heavily in the first principal component (27.4%) are 
width of the outer perianth (WOP), width of the inner perianth (WIP), 
shape of inflorescence (SOI), and plant height (PSH). The second component 
accounts for 25.0% of the total variation and is mostly concerned with 
length of the perianth tube enclosing the ovary (LPO), SOI, PSH, length of 
inflorescence minus flowers (LSI), length of the outer perianth (LOP) and 
length of inner perianth (LIP). The third component accounts for 12.4% of 
the total variation and number of flower per scape (NFS), length of the lowest 
bracts (LLB), and width of the widest leaves (WWL) are highly weighted. 

Results from two-dimensional plots of component 2 against components 
1 and 3 (Figs. 2 and 3) show that the type specimen of // . mkrantha is margin- 
ally included in the continuous range of variation described by the OTUs of 
H. hakuunensis (populations A-N, P and Q) including the type (R). The range 
of variation for H. hakuunensis is distinct from that of//, middendorffii . 


Hemerocallis middendorffii, found on open areas of central and northeastern 
mountains in Korea (S.S. Kang pers. obs.; S.H. Oh pers. comm.), differs from 
H. hakuunensis by having a capitate-type inflorescence; large inflorescence 
bracts (2.0-4.0 cm long, 1.0-3.0 cm wide); short perianth tube (1 .0-1 .8 cm 
long); short scape (30-47 cm long); and a different flowering period from 
middle May to June. The results from principal components analysis 
revealed these two species are distinct from each other (Figs. 2-3). 

Nakai (1943) originally described H. micrantha based on only one 
herbariumspecimencollectedbyO.SyogofromHamyang,Prov. Gyeongsang 
Nam of southern Korea. He noted that H. micrantha is distinct from other 
species by having small-sized orange-yellow flowers and a highly branched 
inflorescence. For example, he reported that the length and width of 
perianth lobes are 4 cm and 5-6 mm, respectively, and the length of 
inflorescence is 23 cm long. These values are included in the continuous 
range of variation observed during our field trips for H. hakuunensis. 
Individuals with 20-30 cm long mflorescences were encountered within the 
populations of D, K, and M. Individuals with small-sized (ca. 4-6 cm long) 
length of perianth lobes were also observed within the populations of A, B, 
D, E, I, M, and N. In addition, the flowering date (17 July) and collection 
location (Prov. Gyeongsang Nam of southern Korea) for H. micrantha fall 
within the range of flowering period (June to July, August in the mountain- 
ous areas) and geographic distribution (southern, central, and northwestern 
Korea) of H. hakuunensis. It is highly probable that the type of H. micrantha 
is one of the variable individuals observed for H. hakuunensis as revealed by 
PCAs (Figs. 2-3). Although H. hakuunensis and H. micrantha were published 
in 1 943 on the same volume oi Journal of Japanese Botany (pp. 3 1 5-3 1 6), H. 
hakuunensis has priority over H. micrantha because the former was described 
ahead of the latter. So H. hakuunensis is the correct name for the taxon. 

Kang and Chunc;, Heme 

FIGS. 2-3. Diagrams showmg the results ofPCA 
hakuunenm and H. mtcrantha. FIG. 2. PCA (compo 
FIG. 3. PCA (components 2 and 3) composed of 8' 

nsis Nakai, J. Jap. Bot. 19:315. 1943. Type: KO- 
am: Mt. Baekun, 22 Aug 1934, Nakaj s.n. (holotype: TI!; 
: KOREA. Prov. Cholla Nam: Mt. Chiri, 25 Jul 1937, 0. 

Syogo . 

Hemerocallis mkrantha Nakai, J. Jap. Bor. 19:315-316. 1943. Type: KOREA. Prov. 
Gyeongsang Nam: Hamyang, 17 Jul 1937, 0. Syogo s.n. (holotype: TI!). 

Herbaceous perennials. Roots usually tuberous (ca. 1.5-2.0 cm long, 0.8 
cm wide), grayish yellow. Leaves 40-100 cm long, 1.2-2.5 cm wide, 
greenish yellow. Scapes ascendant or erect, 32-145 (80 ± 1 5 .62, mean ± SD) 
cm long, nearly as long as or shorter than leaves, usually 1-2 lanceolate 
bracts below the inflorescence, 1.2-6.0 (9-5) cm long, 0.7-1.8 cm wide; 
inflorescence branched (rarely Y-type) with 4-27-flowers, 3-30 (10.82 ± 
4.43) cm long; bracts ovate, greenish, membranous on margin. Perianth 
(fresh) orange-yellow, 7-10 cm long, tube 1.5-3-0 cm long, green with 
orange tint; throat orange yellow; inner perianth lobes 4.5-9.0 cm long, 
1.6-2.6 cm wide. Stamens 3-4 cm long, inserted; filaments orange-yellow, 
attached to the base of the perianth tube; anthers ca. 6 mm long, dark brown 
with purple tint. Style filiform, ca. 8.5 cm long, exerted beyond the stamens. 
Capsule usually oblong-oval, 2.0-2.5 cm long and 1.1-1.5 cm wide, cross- 
wrinkled when dried, surface usually covered with wart-like projections, 
apex emarginate; seeds shining black, angled, 6 mm long, 4 mm wide. 
Flowering from June to July (August in the mountainous areas) in Korea; 
fruit ripening late July to September. 

Korean name: Baekunsan-wonch'uri 

This species is commonly found on the humus or granitic soils and open 
areas or under pine-oak forests on hillsides of southern, central, and 
northwestern Korea, including Kojae, Namhae, Komun, Dolsan, Pogil, 
Chin, Anmyeon, and Kangwha islands (Korean endemic species). 

We thank Jungo Noguchi and Mitsuru Hotta for discussions on the 
classification oi Hemerocallis. Special thanks go to Sun Gi Chung, Duk Seo 
Gu, and Ou Kun Chin for their company on field trips. Also, appreciation 
is extended to Barney Lipscomb and two anonymous reviewers for com- 
ments on the manuscript. We are also grateful to the directors and staffs of 
the National Parks and the Department of National Monuments of Culture 
Property preservation Bureau of Korea for permission to collect samples 
from Hallasan, Hanryohaesang, Sobaeksan, Sokrisan, Tadohaesang National 
Parks and Hong and Cheju islands. We especially thank the herbaria 

directors and curators of GA, KYO, MO, SNU and TI. This research was 
supported in part by a National Science Foundation Dissertation Improve- 
ment Grant (BSR-8914430) to MGC, a Korea Research Foundation Non 
Directed Research Fund (1992) to MGC, and a Korea Science and Engineer- 
ing Foundation Grant (931-0500-031-2) to MGC. 

Bailey, L. 1930. Hemerocallis: The day-lilies. Genetes Herb. 2:143-156. 
Chung, M.G., S.B. Jones, J.L. Hamrick and H.G. Chung. 1991. Morphomerric and 

isozyme analysis of the genus Hosta (Liliaceae) in Korea. Plant Species Biol. 6:55-69. 
Cohen, S. 1986. The well-tinkered daylily. Garden 16:19 (Sep/Oct). 
HoTTA, M. 1986. Hemerocallis auranttaca group in northern Kyushu, Japan. Acta Phytotax. 

Geobot. 37:17-21. (in Japanese) 
KiTAMURA, S., G. MuRATA and T. KoYAMA. 1986. Colored illustrations of herbaceous plants 

of Japan (Monocotyledoneae). Hoikusha Publ. Co., Ltd., Osaka, Japan, (in Japanese) 
Lee, T.B. 1985. Illustrated flora of Korea. Hyangmoonsa, Seoul, (in Korean) 
Matsuoka, M. and M. Hotta. 1966. Classification oi Hemerocallis in Japan and it's vicinity. 

Acta Phytotax. Geobot. 22:25-43. (in Japanese) 
Nakai, T. 1932. Hemerocallis japonica. Bot. Mag. Tokyo 46:111-123. 
Nakai, T. 1943. Notulae ad plantas Asiae Orientalis (XXVIII). J. Jap. Bot. 19:315-316. 
Sas, 1987. SAS/STAT Guide for personal computers, ver. 6ed. SAS Inst. Inc., Cary, NC. 
Sneath, P.H.A. and R.P. Sokal. 1973. Numerical taxonomy: The principles and precise of 

numerical classification. Freeman Co., San Francisco, CA. 
Stout, A.B. 1941, Memorandum on a monograph of the genus Hemerocallis. Herbertia 


International Institute for Sustainable Development (USD). 1992. Sour- 
cebook on Sustainable Development. (ISBN 1-895 5 36-04-9, pbk). 
USD, Winnipeg, Manitoba, Canada R3B 0Y4. $20.00. 133 pp. 

and an extensive bibliography categorized by topics and regions, the sourcebook describes 
audio-visual organizations and materials available as well as a detailed guide to relevant 
computer networks, conferencing systems, and databases. — Larry Ford. 

Banuri, Tariq and Frederique Apffel Marglin (Eds). 1993. Who Will 
Save the Forests?: Knowledge, Power and Environmental De- 
struction. (ISBN 185649-159-5, hbk; 1985649-160-9, pbk). The 
United Nation University, World Institute for Development Econom- 
ics Research, Helsinki, Finland; Zed Books Ltd., London, England. 
$55.00 (hbk); $17.50 (pbk). 195 pp. 




Botanical Survey of India 

DebraDun-248 195, INDIA 

R.R. RAO and A. GARG 

National Botanical Research Institute 
Lucknow-2 26001, INDIA 

A new species of Poaceae, Saccharum stewa, 
species is analyzed and compared with its c 

e inconografia una especie nueva de Poaceae, Sacchan 
jarg, de Jammu y Cachemira, noroeste del Himak 
linologia de esta especie con la de su patiente cetcar 

The genus Saccharum L. (sensu lato), with ca. 3 5-40 species, is distributed 
in the tropics but extends to warm temperate regions of the world. In India, 
the genus has ca. 16 species mostly in the tropical belt (Bor I960). During 
a taxonomic study on grasses of Jammu & Kashmir, the authors came across 
an interesting specimen of the genus collected from the Kashmir Valley and 
carrying a determinavit slip by N.L. Bor bearing the name Erianthus stewartii 
Bor. Critical examination of the specimen indicated that this is an unique 
species morphologically very near to Saccharum griffithii Munro ex Boiss. 
However, the binomial Brianthus stewartii was never published and further, 
as the species of Erianthus are now transferred under Saccharum, a new 
binomial, Saccharum stewartii is necessary and described here. 

Saccharum stewartii Rajeshwari, R.R. Rao & Garg, sp. nov. (Fig. 1). 
Proxime affinisj". griffithii Munro ex Boiss. sed differt racemis perlongioribus, pedunculis 

culms up to 3 m high. Leaf blades up to 50 cm long, tough, 

-36. 1994 

6-7.5 cm long; internodes hairy; hairs up ro 6 mm long. Sessile s; 
to 8 mm long; callus bearded with silky hairs; hairs up to 6.8 
usually shorter than the spikelets and not concealing them; glu 
ovate-lanceolate, keeled, 1-3 nerved, scabrid on the keels, hairy c 
or glabrous; lower lemma 3 mm long, 3-nerved, ciliate at th 
hyaline; lower palea membranous, as long as the lower lemma, ci 


margin; upper palea membranous, hyalme; upper lemma 2 mm long, entire, 
ciliate at the margins, awned; awn 3 mm long, straight; anthers 2 mm long. 
Pedicelled spikelets male only or bisexual; lodicules 2, cuneate or ovate- 
oblong, faintly ciliate; caryopsis not observed. 

Type: INDIA. Kashmir: Rupal to Gunkoc, 2500 m, 23 Aug 1939, R.R. Stewart & l.D. 
Stewart 18893 (holotype: DD). 

Saccharumstewartii closely resembles S. griffithii Munro ex Boiss in having 
awned spikelets in short, articulate racemes borne on long panicles, acute or 
acuminate, membranous glumes, and lower florets all neuter. The two 
species can be separated as follows: 

hairs yellow or cream colored, 3-4 mm long; awn of upper lemma 4 mm 

long S.gnffithtt 


In addition to the above morphological differences the new species also 
reveals certain palynological differences with that of S. griffithii. The 
morphology of the pollen grains in the two species is discussed below. 

Saccharum stewartii Rajeshwari, R.R. Rao & Garg 

Grains 29-3 pm (29-29.6 pm) in diam, spheroidal with circular outline; 
typically monoporate, pores opercuiate, annulate. Annulus conspicuous. 
Exine surface finely granulate, granules of low relief, grouped in close 

(2.25 pm), (Figs. 2 & 3). 
Saccharum griffithii Munro ex Boiss 

Grains 16 pm (15-18 pm) in diam, spheroidal with circular outline, 
typically monoporate, pores opercuiate, annulate. Annulus faint. Exine 
surface psilate with few punctae distantly scattered having a negative 
reticulum. Exine 1 pm thick, slightly varying in thickness near the aperture 
(Figs. 4 & 5). 

The authors are thankful to the Director of the Nati 
Research Institute, Lucknow for facilities and encouragerr 
S.K. Jain, Emeritus Scientist, National Botanical Researc 
going through the manuscript. One of us (SR) is thankful to 
the Botanical Survey of India, Calcutta for the award of a S 
Fellowship during the tenure of which this work was carrie 






The Wilderness Society 

900 Seventeenth Street NW 

Washington, DC 20006, U.S.A. 


Department of Forest Sciences 

Colorado State University 

Port Collins, CO 80523, U.S.A. 


Center for Ecological Management of Military Lands 

Colorado State University 

Fort Collins, CO 80523, U.S.A. 

n alahamenm E. A. Smith ex Chapman, a Cate^ 
d for addition to the List of Endangered and Thre; 
km from any previously known population. This 

Una poblacion alopatnca de Croton alabamemis E. A. Smith ex Chapman, Categon'a 2 c 
Especie Candidata, que esta siendo considerada para ser agregada a la Lista de Plantas e 
Peligro de Extmcion, fue descubier'a a una distancia de 1000 km de la poblacic 
previamente conocida. En este tra^ ajo se describe la extension y estado de la poblacic 
alopatrica. La poblacion de Croton esta situada a lo largo del fondo de un caiion aparentanc 
poder sobrevivir y crecer en buenas condiciones. La ocurrencia de Croton no parece ten( 


Croton alabamemis E, A. Smith ex Chapman (Euphorbiaceae) has been de- 
scribed as "one of the rarest shrubs m the United States" (Farmer and Thomas 
1969). It is a short-lived (<20 y), multi-stemmed, monoecious shrub <3 m 
tall found primarily on limestone and shale outcrops along the Warrior and 
Cahaba Rivers, Tuscaloosa and Bibb counties, Alabama. It was collected 
once in 1899 in Tullahoma, Coffee County, Tennessee, but has not been 
reported from there subsequently. Croton alabamemis has long been thought 
to grow only in isolated populations within these two neighboring counties. 
Due to its restricted range, it is being considered for addition to the List of 
Endangered and Threatened Plants under the Endangered Species Act of 
1973, as amended. It currently is designated as a Category 2 Candidate, 
which means that "there is some evidence for vulnerability, but . . . there are 
not enough data to support listing proposals at this time" (USDI 1991). 

In early 1990, Croton alahamenm was discovered over 1000 km from any 
previously known population on the U.S. Army's Fort Hood, Texas. The 
population was discovered by John Cornelius, a wildlife biologist with the 
Fort Hood Resource Management Department, during an excursion to view 
bird habitat in the Owl Creek Mountains. Cornelius showed the population 
several weeks later to Carol Beardmore of the U.S. Fish and Wildlife Service 
and Rex Wahl of Texas Parks and Wildlife. Ginzbarg (1992) mistakenly 
credited the discovery to Beardmore and Wahl. Not long afterwards, a 
second and a third population were discovered in Travis County, to the 
south. The species identity was determined by Steve Ginzbarg, a graduate 
student in botany at the University of Texas, Austin. Ginzbarg (1992) 
subsequently determined that the Texas plants are sufficiently distinct to 
merit varietal status. He consequently described them as Croton alabamemis 
E. A. Smith ex Chapman var. texemis Ginzbarg (Ginzbarg 1992). Key 
characteristics of the Texas variety are represented in Figures 1 and 2. 

In June 1991, we undertook to describe the extent and status of the 
population at Fort Hood as part of the Department of Defense's Legacy 
Resource Management Program. The Legacy Program was established to 

military lands. Our study is aimed at gathering the requisite information to 
effect proper stewardship of this species at Ft. Hood. We therefore restricted 
our analysis to only one of the three Texas populations (i.e. the Ft. Hood 
population). We recommend that similar studies be undertaken in both 

FIG. 1. Croton alahamensts var. texei 
rimary and secondary branches (flow( 
lowing fully developed leaves. 

Texas and Alabama to assess the status of all populatii 
scientifically important species. 

This study was conducted in several small watersheds of the Owl Creek 
Mountains, Bell and Coryell counties, in the Hill Country of central Texas 
(31°N, 97°W). The Owl Creek Mountains (elevation ~300 m) are com- 
posed of Mesozoic limestone overlain by clayey and loamy soils (McCaleb 
1985). The formation rises to its summit plateau -65 m above the west-to- 
east flowing Owl Creek in less than 2 km . Intermittent tributary streams run 
northward to the creeksuch that virtually all canyon walls have east- and 
west-facing aspects. 

The climate is hot in the summer, and the winters are generally mild, with 
an occasional cold surge. The average daily temperature during the summer 
is 28°C, with an average daily maximum of 36°C. Winter daily tempera- 
tures average 9°C, with an average low of 3°C. Rainfall is distributed uni- 
formly throughout the year, with a slight peak in the spring and an average 
annual total of 825 mm (McCaleb 1985). Owl Creek flows to some extent 
throughout the year, but the tributary streams are dry most of the year. 

The vegetation of the area (see Appendix 1 for complete plant species list) 
is primarily Ashe juniper (Juniperus ashei Buchh.) woodland of the appropriate 
character to meet the habitat needs of two rare birds, the black-capped vireo 
{Vireo atrkapilla) and the golden-cheeked warbler {Dendroka chrysoparia). 
The vireo primarily inhabits the shrubby balds of the summit plateau, but 
the warbler requires the bark of relatively old junipers for nesting materials 
and mature hardwoods for feeding (J. Cornelius pers. comm.). 

Prior to our study, Croton alabamemis var. texensis had been identified from 
two of the tributary canyons of Owl Creek. We searched five additional 
canyons and found only three more plants in the canyon between the original 
two and several plants along the stretch of Owl Creek between the two 
tributaries; populations oi^ Croton appear to be largely restricted to the two 
original canyons. Our study involved two phases: a description of the 
structure and habitat of the two Croton populations (canyons 1 and 3) and an 
attempt to discern why Croton is all but absent from the canyon between 
(canyon 2). Subsequent to the completion of this study, another population 
consisting of 35 individuals was located in a canyon three km east of the 
study area. This population is not considered in our analysis. 

The bed of each of the three canyons was mapped using a tape and a hand- 
eld compass. The slope of the creek was measured using a hand-held 
iinometer. Three high density stands of C. alabamensis var. texensis were 

identified for intensive sampling along the creekbeds in Canyons 1 and 3. 
At each sample location, two transects were established perpendicular to the 
creekbed to facilitate sampling of both east- and west-facing aspects. In 
Canyon 2, which lacked Croton, similar pairs of transects were established at 
distances up the canyon comparable to those identified in Canyons 1 and 3. 

Slope topography, overstory and understory cover, Croton seedling den- 
sity, Croton adult population structure, and soil depth were determined 
along each transect. At each site, a tape was stretched upslope beyond the 
extent of the Croton population. In some cases, this was as far as 60 m from 
the creekbed, but was generally less. Transects in Canyon 2 were 30-40 m 
long. Slope breaks and important topographic features were noted along the 
slope, and a cross-section map was prepared for each transect. Soil depths 
were measured at 10 m intervals and at important topographic features by 
probing the soil with aim rod. 

Along the tape, cover of overstory (>2 m tall) and understory (<2 m but 
>10 cm tall) vegetation was determined using the line-intercept method 
(iVlueller-Dombois and EUenberg 1974). The line also served as the center 
of a 2 m wide Croton seedling belt in which individuals <30 cm tall were 

Croton >30 cm tall were sampled using a modified nearest-neighbor 
method adapted from those described by Muelier-Dombois and EUenberg 
(1974). Starting at the beginning of the line (middle of the creekbed), a 1 80° 
arc was searched upslope for an individual of Croton >30 cm tall. The 
distance and azimuth to the nearest plant were recorded as were the height 
of the tallest shoot and the diameter of each live shoot (to the nearest 0.5 cm). 
From that plant, the process was repeated until no plants could be found 
within 10 m of the last plant (Fig 3). In only one case, the process was 
suspended and moved back to the tape when the search led to a plant >10 
m from the tape. Thus, all plants were sampled within 10 m of the tape. 

Consistent with other nearest-neighbor methods, the distance between 
plants was assumed to be related to the share of the total area allotted to an indi- 
vidual plant. From these data, population density could be calculated as the 
total number of plants sampled divided by the sum of areas allotted to indi- 
vidual plants. Additionally, using a simple trigonometric conversion, the den- 
sity represented by individual plants could be plotted against distance from 
the creek to assess changes in density related to environmental variables. 

Sizes of individual, multi-stemmed Croton plants were described using the 
statistic of equivalent diameter (Dgq). Equivalent diameter is the diameter 
of a single stem that possesses the same cross-sectional area as the sum of all 
the stems borne by the plant. For example, as the total cross-sectional area 
of four shoots, each 2 cm in diameter (4 X 3.14 cm^) is equal to that of one 
shoot, 4 cm in diameter (12.56 cm^), the Deq for the four shoots is 4 cm. 

Ant ecology 

The population structures of Croton alabamensis var. texensts in the two 
canyons are presented in Figure 4. In both canyons, Croton was well 
represented in the smaller diameter classes, but also occurred as large mature 
plants. This decreasing monotonic population structure was found in all 
transects in which Croton was relatively dense. Therefore, both canyons 
appeared to support healthy populations of adults, juveniles, and new 

Croton density was highly variable within Canyons 1 and 3. Each canyon 
contained places in which scarcely an individual was found. In other places, 
Croton formed dense thickets in which it dominated to the near exclusion of 
other understory species. The densest stands occurred in Canyon 1 where 
Croton density (adults and seedlings) in two transects exceeded 100 plants/ 
100 m^, but three transects in Canyon 3 exceeded 50 plants/100 m^ (Table 
1). Overall, we estimate the number of plants occurring in the 1.5 km of 
creekbed in Canyons 1 and 3 to be ~ 20,000 individuals. 

In general, Croton was found only in the canyon bottoms. No individuals 

s found ; 

1 from the c 


nly four contained plants >40 m from the creek. Croton density fluctuated, 
ut Croton did not decline proportionally with distance from the creek. 

Canopy gaps along the line intercept showed no correlation with Croton 
ccurrence. Croton were found in openings and in the deepest shade. The can- 
ons of the Owl Creek Mountains have been subjected to harvesting of juni- 
er trees for fence posts, and some of the gaps may have been anthropogenic. 

Canyon 1 

10 12 14 >15 

4 6 8 10 12 14 >15 

Deq Class (nnm) 


2 ^ '^^^ 






3 2 84 





2'i '4I 

3 3 96 





9.7 55.8 

3 5 99 





23.4 28.6 

In some cases, the loggers incidentally cut adult Croton along roads and skid 
trails, but the plants sprouted and were growing well. 

Topographic features also showed no direct correlation with Croton 
density. Croton occurred on bank slopes, terraces, and on toeslopes of each 
canyon. Soil depth, however, explained much of the variability in Croton 
distribution. A comparison of soil depth with the presence of Croton along 
the transect indicated a significant association of adult plants with deep soil 
(p<0.05; Kruskal-Wallis test). Our inability to measure depths in excess of 
1 m means that the measured mean soil depth in the presence of Croton (0.80 
m; n = 64) was probably considerably less than the true mean. In contrast, soil 
measurements in the absence of Croton averaged only 0.42 m (n = 33) in 
Canyons 1 and 3. 

Comparison of Croton density with overstory species composition sug- 
gests an association with mesic sites, as would be expected from the observed 
restriction of Croton to the canyon bottoms. Figure 5 shows the relationship 
of adult Croton density to the combined cover of Fraxinus texensis (Gray) Sarg. 
and Quercus muhlenbergii Engelm. and to the combined cover of J uniperusashei 
and Quercus texana Buckl. within 40 m of the creek. These four species 
dominated the overstory stratum in these canyons; only the woody vine Vitis 
mustangensis Buckl. contributed comparable cover over the study area. Of the 

— 0.2 0.4 0.6 0.8 1.0 

^ Juniperus and 0. texana Cover 

0.0 0.1 0.2 0.3 0.4 0.5 

Fraxinus and Q. muhlenbergii Cover 

three species of oak encountered during sampling, Q. muhlenbergii is the most 
mesic (Miller and Lamb 1985), and Fraxinus is found primarily in canyons 
(Correll and Johnston I910).juniperus is not so restricted, and Q. texana is 
described as occurring on "dry limestone hills and ridges" (Miller and Lamb 
1985). In the study area, Croton was more abundant on mesic transects in 
which Fraxinus and Q. muhlenbergii were common and less abundant on dry 
transects in which J. ashet and Q. texana were most common. 


The four tree species listed above dominated the vegetation of all three 
canyons. Figure 6 shows the contribution of these species to the overstory 
cover within 40 m of the creek (40 m was used as the limit oiCroton habitat). 
The dominant species in all three canyons was clearly J. ashei, which con- 
tributed about one-third of the cover. The other three species collectively con- 
tributed another third. These four were the only species to contribute > 10% 
relative cover in one of the three canyons, although some species, such as V 
mustangensis, Ulmus crassifolia, Nntt. Juglans major (Torr.) Heller., and Celtis 
laevigata Willd. were abundant locally (Appendix 2). All overstory species 
contributing < 1 0% relative cover are included in Figure 6 as "minor species. " 

canyons supporting Croton populations and Canyon 2 , we subj ected transect 
cover data to principal components analysis (PCA) (SAS/STAT User's Guide 
1988). If the transects in Canyon 2 differed in overstory composition from 
the other two canyons, those sites would have segregated as a distinct habitat 
type. Canyon 2 showed no difference in habitat from the other two as 
expressed in overstory composition (Fig. 7). 

Likewise, understory cover displayed no pattern related to Croton occur- 
rence. With the exception of Croton itself, all three canyons supported 
similar understory communities (Appendix 2). In the two canyons in which 
it occurred, understory Croton cover averaged 10.4%. (Again, placement of 
the transects was biased by the presence of dense populations.) Other 
relatively abundant understory species included Fraxinus species (5.5%), V 
toxicodendron L. (2.6%), various grasses (2.4%), Q. texana (2.1%), and Ilex 
decidua Walt. (2.0%). 

The three canyons differed very little in both total overstory and under- 
story cover. Overstory cover ranged from 50% in one transect in Canyon 1 
to 99% in two transects in Canyon 3, but no significant difference occurred 
among canyons (Table 1). Similarly, understory cover ranged from 16% in 
one Canyon 2 transect to 54% in a Canyon 3 transect, but canyon means 

Canyon 1 Canyon 2 Canyon 3 

I I Juniperus ashei 

■I Quercus texana 

■ Fraxinus texensis 

■ Quercus muhlenbergii 
I I Minor species 


Steepness of canyon walls within 40 m of the creek varied widely from site 
to site (10% to 60%), but means from the three canyons were similar (Table 
1). Canyons 1 and 3 were the largest watersheds, but watershed areas above 
Croton populations in Canyon 3 were no larger than those present in Canyon 
2. Likewise, soils of a depth that favored Croton in Canyons 1 and 3 were 
abundant in Canyon 2. The only morphological feature distinguishing 
Canyon 2 from Canyons 1 and 3 was the steepness of the creekbed. 
Streambed gradient in Canyon 2 (3.81%) was significantly steeper than in 
Canyons 1 and 3 (1.92% and 2.11%) (p<0.05; Kruskall-WaUis test). 


Croton alabamenm 

var. texem 

m in the O 

wl Cree 

'k Mountains grows in 

healthy, self-sustainii 

ng popular 

ions along tl 

tie botto 

rns of tributary canyons 

and the connecting 

section of Owl Creek. 


occurrence exhibits no 

association with ovei 

-story gaps 

;, disturbanc 

:e, or particular fluvial geomor- 

phic features. It appears to be 

restricted tc 

. canyor 

1 bottoms characterized 

only by mesic condii 

tions provi 

ded by the ] 

presence of overstory cover and 

deep soils. There is : 

some suggestion that 

high CO 

ver of R texerms and Q. 

muhlenbergii indicate: 

s a good SI 

te for Croton 



33 1 2 


3 1 

1 1 

liffer in c 

The distribution oiCroton does not appear to be limited by the availabihty 
of sites. Canyon 2, which contained only 3 plants, supported a similar 
overstory and understory to Canyons 1 and 3. Canyon 2 also contained deep 
soils and a similar geomorphology to the other two canyons. Likewise, 
watershed size did not appear to explain Croton presence. The only feature 
that differed among canyons was stream gradient. 

Hupp (1988) discusses several ways in which stream channel geomor- 
phology affects vegetation, including fluvial landforms, flood frequency and 
duration, and stream gradient. It would be consistent with other observa- 
tions if the steeper gradient of Canyon 2 afforded a less mesic environment 
that discouraged Croton survival. However, fluvial processes in these arid 
intermittent channels are quite different from those of the eastern floodplain 
systems that Hupp discusses, and channel gradient is offered only specula- 
tively as a factor explaining Croton distribution. 

The larger question of how C. alabamemu var. texemis came to occupy 
disjunct sites in Alabama and Texas is beyond the scope of this study. 
Although Ginzbarg (1992) discusses the possibility that the present distri- 
bution may be an ice age relict, he suggests that the Texas populations may 
be more easily explained as the result of relatively recent introduction by 

long distance seed dispersal by birds. Regardless, it is likely that the disjunct 
distribution resulted from a prolonged process of migration, colonization, 
and extinction of many sites. The absence of Croton from Canyon 2 may 
reflect not the lack of available habitat, but the vagaries of colonization and 
extinction. Perhaps a population once thrived there and has gone extinct, or 
seeds have, by chance, not recently reached this canyon. More likely, the 
canyons of the Owl Creek Mountains and similar features of the Edwards 
Plateau support metapopulations of Croton alabamensis var. texensis, each 
subpopulation establishing, thriving, and going extinct only to be replaced 
somewhere else. Only through repeated observations over long time periods 
will we understand the process. 

Regardless of the limitations on its range within the study area, the 
behavior of C. alabamensis in the Owl Creek Mountains is dramatically 
different from its behavior in Alabama. Farmer (1962) describes Croton as 
occurring on shallow soils and rock outcrops at mid-slope positions in two 
counties in Alabama. He describes the habitat as "shallow soil ... on 
moderately- to steeply-sloping terrain; high temperature of soil and air 
during summer; intense drought; and freedom from fire." Soil is described 
as "usually only a few cm thick and seldom more than 1 m thick." In our 
study site, Croton occurs on deep soils on toeslopes and fluvial deposits of 
canyon bottoms. 

The groves in Alabama are "marked by shrub dominance, few or no large 
trees, and a relative absence of herbs" (Farmer 1962). Occasional plants are 
found under the forest canopy surrounding the outcrops, but these are 
considered only "extensions of nearby thickets". This contrasts sharply with 
Croton behavior in Texas. In our study sites, Croton occurs as an understory 
shrub in the company of many large trees and a healthy herb layer. 

Farmer considers adaptation to extreme drought to be an important factor 
in the ecology and distribution of C. alabamensis in Alabama (see above). In 
Texas, however, this species behaves as a drought avoider by remaining in 
mesic canyon bottoms. Perhaps these differences result only from perceptions 
relative to annual climate and surrounding vegetation. The 3-5 inches (7-12 
cm) of monthly precipitation that Farmer ( 1 962) reports for the "dry" season 
in Alabama is considerably greater that the 2-3 inches (5-7 cm) recorded for 
summer months in central Texas (McCaleb 1985). In Texas, where upland 
vegetation is necessarily drought adapted, a species requiring 7-12 cm of 
precipitation naturally would be restricted to the most mesic sites. 

Another interesting difference between Croton behavior in Alabama and 
Texas regards vegetative reproduction. Farmer (1962) discounts asexual 
reproduction in Croton by stating that "there are no rhizomes or adventitious 
rootings that result in plant reproduction." In Texas, however, we observed 

numerous plants that had produced "new" upright shoots through the nodal 
rooting (layering) of prostrate branches. We did not investigate the degree 
of connectivity between the layered offspring and the parent plants, but 
layering appears to be a potential mechanism of asexual reproduction in C. 
alahamemis var. texensis. 

The disparate behaviors exhibited by this species over its disjunct range 
underscore the necessity for site-specific studies prior to making manage- 
ment recommendations. The ecology oiCroton in Alabama would suggest a 
conservation strategy that might well be unsuccessful in Texas. The Ala- 
bama ecology implies preservation of, and population augmentation into, 
dry, open, limestone outcrops. This strategy, if adopted in Texas, likely would 
fail miserably. 

We know very little of the disturbance ecology of C. alahamensis var. 
texensis in central Texas. The species apparently tolerates some degree of 
physical disturbance* as plants injured or cut off during the pole cutting 
operation appear to have recovered well. We know nothing, however, of the 
species' response to fire or soil disturbance. In Alabama, fire is believed to be 
lethal to Croton (Farmer 1962). We saw nothing to indicate a dependence on 
disturbance for establishment. Conservation efforts, therefore, should focus 
on maintenance of the undisturbed nature of the mesic forests of the area. 
Construction of new roads for military training and logging should be 
discouraged, and old roads should be allowed to fall into disuse. The 
importance of this plant community has already been recognized for a rare 
bird; it now appears to be critical to the survival of a rare plant as well. 

We thank Dennis Herbert, Billie Ray Jones and John Cornelius, Division 
of Wildlife, Fort Hood, Texas for their assistance and encouragement during 
the study. A special thanks to Keith Schulz for field assistance and to Tracy 
Wager for the line drawings. Funding was provided by the USDA Forest 
Service, Rocky Mountain Forest and Range Experiment Station, Department 
of Defense Legacy Resource Management Program, and the Natural and 
Cultural Resources Division, Engineering and Housing Support Center. 








Calyhphus drumrm 


Smilax bona-nox 




Sophora secundiflo 






ULCR 2.02 










CoRRELL, D.S. and M.C. Johnston. 1970. Manual of the vascular plants ol Texa: 

Research Foundation, Rennet, TX. 
Farmer, J.A. 1962. An ecological life histoty oiCroton alahamensis E.A. Smith ex Ch 

Unpublished Ph.D. Dissertation, University of Alabama. 
Farmer, J.A. and J.L. Thomas. 1969. Disjunction and endemism in Croton alab 

Rhodora 71:94-103. 
GiNZBARCi,S. 1992. A new disjunct variety ofCro/o«^/,//;.rfWOTm(Euphorbiaceae)fron 


Miller, H.A. and S.H. Lamb. 1985. Oaks of No 

Happy Camp, CA. 
McCaleb, N.L. 1985. Soil survey of Coryell Cc 

Agriculture. Soil Conservation Service. U.S. 
MuELLER-DoMBOis, D. and H. Ellenberg. 1974 

John Wiley and Sons, Inc. New York. 
Sas/Stat User's Guide. Release 6.03 Edition. ^ 

:h Amei 



hers. Inc. 

mty, Texas. Unitec 
xovernment Printi 
Aims and method; 

i States 
ng Offi. 


tment of 

^8. SAS 


Inc. Cat 



and pla. 



: taxa f 

3r listing 


Smith, Nigel J.H., J.T. Williams, Donald L. Plucknett, and Jennifer P. 

Talbot. 1992. Tropical Forests and their Crops. (ISBN 0-8014- 

277 1-1, pbk). Comscock Publishing Associates, a Division of Cornell 

University Press, Ithaca, NY. $27.95 (pbk), $69.95 (hbk). 568 pp. 

"Many cultivated planes important for food and income in developing countries arose in 

tropical forests." A number of these plants are the subjects of this book, grouped into 

"Beverage and confectionery crops" (e.g., coffee, cacao); "Major fruits" (e.g., mango, citrus, 

pineapple); "Regional fruits" (e.g., durian, rambutan); "Rubber, oils, and resins" (e.g., oil 

palm, tropical pines); "Daily bread" (e.g., bananas, breadfruit); "Fuelwood, fodder, and 

woody grasses" (leucaena and bamboo); "Spices and natural food colorants" (e.g., clove, 

these crops are history, evolution, domestication, spread, genetic resources, gene pools, gene 
crop. Additional chapters discuss the search for new plants to domesticate and the need for 


5 of cons 

ervation. Most entries ir 


ge list of "References" 

(largely from 

the 198 


ack-and-white photo; 

drawings anc 

1 is well 

indexed.— >/.« W. Tk 


National Reseaf 

LCH Council. 1993. 


Grass. A Thin ( 

against Erosion. (ISBN 0-309-04269 

-0, pbk). Nations 



ington, DC. No pr. 

ice given 


Vetiver, V. 

ntveria : 

zizaniotdes, has long bei 

;n cultivat 

:ed as the source of e 

vetiver used i 


s also a gn 

iss of much promise f 

soil erosion ir 

treas is being increasing! 

Hedges of this deep-t 

densely cespi 


ms" strong enough to 

Although its 

hedge plant IS not new, 

it has rece 



United States Department of Agriculture 

Systematic Botany & Mycology Laboratory 

Bldg 003, Rm. 253, BARC-West 

Beltsville, MD 20705, USA 


Department of Botany 

National Museum of Natural History 

Smithsonian Institution 

Washington, DC 20360, USA 


Fort Collins, CO 80326, USA 

Recent c 

ollections fi 

-om Ecuador have uncovered the preset 

ice of a species, Ermh 




ch little comparative taxonomic informati 

ion is available. Previou 


reported on 

ly from Pert 

1, this species is characterized by features 

umque within the gen 


ical description is presented and based . 

3n studies of all availal 



ps with other species are discussed and based on the presence 


shared taxoi 

aomically ir 

nportant characters. Diagnostic features 

of£. weberbauen mcYud 

muticQus ar 

id smooth u] 

pper floret, perennial habit, paired spikel 

ets,spikelets 3.1-3.9 n 


long, and rl 

.e absence o 

fa first glume. 



:es en Ecuador han descubierto la presenci 



Mez, de la q 

ue hay disponible muy poca informacion 


iamente solo 

. de Peru, esta expecie presenta unas caract 

eristicas unicas dentro c 



descripcion morfologica completa en ba 

Lse a estudios de todos 


i disponible: 

5. Se discuten las relaciones con otras especies en base a la presen. 


de caractere 

s compartid( 

OS taxonomicamente importantes. Los cat 


weberbauen j 

incluyen una flor superior mutica y lisa, habito pere: 

nne, espiguillas paread 


espiguilks : 


, de largas y la ausencia de la primera gk 


The Pa 

niceae is a 

. tribe of grasses characterized by 

a paniculate inflore 

cence, spi 

kelets wit 

h two florets, the lower floret ste 

rile, the upper flor 

fertile and coarser in texture than the lower floret. Erwchloa Kunth, a mem- 
ber of this tribe, consists of about 30 species and occurs throughout the 
tropical, subtropical, and warm temperate regions of the world. Presence of 
a cup-like callus at the spikelet base differentiates Eriochloa from the other 
105 genera of the Paniceae. Additional significant characters used to differen- 
tiate Eriochloa within the tribe include an indurate upper floret, disarticula- 
tion at the spikelet base, inflorescence with racemose primary branches, and 
the absence of a bristle (Webster & Valdez 1988, Webster et al. 1989). 
Significant characters used to differentiate the species o^ Eriochloa include 
spikelet shape, presence of an awn on the upper floret, spikelet length, 
presence of hairs in the inflorescence, spikelet arrangement, longevity, and 
the relative length of the florets (Webster 1987, Shaw & Webster 1987). 

Recent collections by Peterson and others in Ecuador have revealed the 
presence of specimens which do not match the morphological characteristics 
of taxa commonly associated with that area. The characteristics of this taxon 
were recorded in DELTA format (Dallwitz 1980, Dallwitz et al. 1993) and 
compared against all other recognized species of Eriochloa. This analysis 
indicated these specimens could not be differentiated from a poorly know 
Peruvian species, E. weberbaueri Mez. Hitchcock (1927) states that this 
species was known only from the type specimen. Tovar (1993) list only two 
specimens, the type and Anderson 740, which was collected in the same area 
of Peru. A survey of collections at the U.S. National Herbarium revealed the 
presence of three specimens {Anderson 740, Sanchez Vega 1031, Camp E-2346) 
which possess the diagnostic characteristics of £. weberbaueri. In addition, G. 
Davidse has identified two MO collections {Sanchez etal. 3367 & Smith 6190) 
of the same species. 

Diagnostic features of Eriochloa weberbaueri include a muticous upper 
floret, a smooth upper floret, paired spikelets on the primary inflorescence 
branches, perennial habit, spikelet length (3.1-3-9 mm long), first glume 
absent, and the absence of elongate rhizomes . Eriochloa distachya, E. michauxii, 
E. polystacyha, E. sericea, and E. weberbaueri are the only species with a 
muticous upper floret and perennial habit. Of this subgroup, E. distachya and 
E. weberbaueri are the only species with long hairs associated with the pedicel. 
Eriochloa michauxii, E. polystacyha, and E. weberbaueri share the presence of 
paired spikelets. Therefore, the presence of hairs at the pedicel apex and 
paired spikelets distinguishes E. weberbaueri within this subgroup of mor- 
phologically related species. Perhaps the most diagnostic characteristic of £, 
weberbaueri is the smooth and relatively shiny upper floret. This characteris- 
tic appears to be unique for E. iveberbaueri within the genus. This species 
appears to be associated with drier environments at elevations between 
1,500 and 3,000 meters. 

fragments at US!). 

Plants perennial; stoloniferous; rhizomatous. Rhizomes with compacted 
nodes; with glabrous scales. Flowering culms 3-8 dm tall; not caespitose; 
erect from the base; not rooting at the lower nodes; with a hard knotty base; 
terminated by a solitary inflorescence; branched from the base; 2-5-noded. 
Nodes not swollen; pubescent. Internodes hollow; glabrous; smooth; neither 
viscid nor glaucous. Leaves cauline or basal; green; without auricles. Sheaths 
6-1 3 cm long; overlapping or not overlapping; not inflated; smooth; glabrous; 
rounded; not keeled; not ciliate; closed. Ligule a fringe of hairs; 0.5-1.2 mm 
long. Collar not differentiated; pubescent. Leaf blades 7-20 cm long; 1.5- 
5 mm wide; linear; flat, involute, or convolute; flexuous; spreading; lax; 
smooth on the lower and upper surfaces; glabrous on the lower surface; 
glabrous or hairy (minutely pubescent) on the upper surface; with a truncate 
base; with flat margins; with margins not thickened; with scabrous margins; 
with glabrous margins; with the midrib not obviously differentiated; 
lacking obvious transverse veins; acuminate. Prophyllum with scabrous 
nerves; with pronounced keels. Peduncle 160-300 mm long; glabrous. 

Panicle 10-35 mm wide; fully exserted at maturity; oblong. Lowermost 
inflorescence node not differentiated; hairy. Main axis present; 60-150 mm 
long; straight; stout; hairy; pubescent; with hairy axils; with quaquaversal 
primary branches. Primary branches 10-30 mm long; 0.3-0.5 mm wide; 
appressed to the main axis or spreading; with appressed secondary branches; 
with secund spikelets; not whorled; 8-13 on the main axis; straight; not 
winged; smooth; hairy; setose; ciliate; with loosely arranged spikelets. 
Rachis terminating in a spikelet. Pedicels present; 0.2-0.5 mm long; 
straight; scabrous; hairy; setose; with hairy apices; discoid at the apex; 
perpendicular with the spikelet base; concave. Disarticulation at the spike- 
let base. Callus differentiated; not prolonged into a stipe; not flared to form 
a discoid receptacle; not oblique; rounded; smooth; glabrous. Cleistoga- 
mous inflorescence absent. 

Spikelets 3.1-3.9 mm long; 0.9-1.1 mm wide; paired; densely overlap- 
ping; evenly distributed on the rachis; not divergent from the axis; homo- 
morphic; adaxial; green; dorsiventrally compressed; lanceolate. First glume 
absent. Rachilla pronounced below the second glume; not pronounced 
between the florets. Second glume 3.0-3.7 mm long; 1 times spikelet 
length; 1 times the length of the lower floret; lanceolate; membranous; 
rounded on the back; smooth; 5-nerved; hairy; pubescent; acute to acumi- 
nate; muticous. Fertile florets 1. Sterile florets 1. Lower floret lacking 

stamens. Lemmaof lower floret 2.8-3.7 mm long; lanceolate; membranous; 
smooth; with equal internerve spacing; 3-nerved; the nerves pronounced but 
not swollen; the nerves smooth; not keeled; with glabrous margins; lacking 
a central longitudinal groove; acute; muticous; hairy. Lower lemma hairs not 
forming a distinct horizontal line; shorter than the upper floret; pubescent; 
smooth and terete; without apical modifications; white. Palea of lower floret 
absent. Upper floret 0.9 times the length of the lower floret. Lemma of upper 
floret 2.3-2.8 mm long; lanceolate to ovate; cartilaginous; slightly convex; 
smooth; yellow; shiny; with involute margins; with margins of the same 
texture as the body; glabrous; 3-nerved; the nerves not pronounced; with 
smooth nerves; without basal modifications; not differentiated at the apex; 
entire; acute; muticous. Germination flap conspicuous. Palea of upper floret 
well-developed; enclosed at the apex; cartilaginous; smooth; the base neither 
swollen nor protruding; the nerves pronounced but not winged. Lodicules 
about 0.1 5 mm long; the nerves not pronounced. Stamens 3. Anthers L5-1.7 
mm long; yellow. Stigmas yellow to orange. South America: Ecuador & Peru. 

Specimens Ex 

amineci: SOUTH AMERICA. Ecuador. Provir 



n American H 

wy at the bridge crossing the Rio Leon, elev. 1880 m, 24 Aj 

1990, P.M. Pe^et 

■son, C.R. Anm 



CO Loja, crossing 

of Rio Leon,. 

;lev. 1800 m, 30 May 1990, P./W.. 

Peterson and E.J. Judziewi. 

9378 (US!); vail. 


ute, between Paute and Cuenca, 2 

6 Apr 1945, Camp E-2 34 

(US!). Peru. Sar 

ua Cruz, /. Sa. 

«f^.z V.i« / 03 i (US!); 11 Apr 19 

48, Anderson 740 (US!). 

Gratitude i 

IS extended i 

to Dr. Gerrit Davidse for ma 

king valuable sugges 

tions concern 

mg the taxc 

)nomy of this group. 

ripcions.Taxon 29:41-46. 
de to the DELTA system: 
1 ed. CSIRO Division ol 

, A.S. 1927. The 

HAW, R.B. and R.D. Webster. 1987. The genus Eriochloa (Poaceae: Panic 

Central America. Sida 12:165-207. 
■qvar, O. 1993. Eriochloa in las gramineas (Poaceae) del Peru. Ruizia 1: 
Webster, R.D. 1987. The Australian Paniceae (Poaceae). J. Cramer, Ber 
and J. Valdes Reyna. 1988. Genera of Mesoamerican P 

, J.H. Kirkbride and J. Valdes Reyna. 1989. New Woi 


QuFROL, Daniel. Undated. 1992. Genetic Resources: A Practical Guide 

to Their Conservation. (ISBN 1-85649-203-6, hbk; 1-85649-204- 

4, pbk). Zed Books Ltd., London, England. $55.00 (hbk); $22.50 

(pbk). 252 pp 

Daniel Querol originally compiled this book as a manual for his course on genetic 

resources and genebanks at the Chapingo Autonomous University in Mexico. He later 

Genetic Resources Program in Peru. Those experiences, over a decade, are reflected in the 

ing, maintaining, and using a genetic resource conservation program. — Larry Ford. 

ACTA BOTANICA HUNGARICA. Volume 36, numbers 1-4, 1990-91. Twenty 
papers. El genero Moacroton Croiz. (Euphorbiaceae). A. Borhidi. — Taxonomic revision of 
genus Leucocroton (Euphorbiaceae) A. Borhidi. — New spontaneous taxa of the genus 
Clerodendrum Brown ex L. (Verbenaceae) in Cuba. 2. Kereszty.— Xylotomkal examina- 
tion of lignifying shoots and roots for age determination ofgrasslands — Fumana procumbens 

species of tree belonging to the Caesalpiniaceae, Fabaceae and Mimosaceae families — 
Caesalpmiaceae II. K. Babos, L.J.C. Cumana.— Xylotomical examinations of some Venezu- 

Caesalpiniaceae III. K. Babos, L.J.C. Cumana.— Comparison of the pollen of various 

G. PALFi-DEiM.^Cunninghamia R. Br. in the pollen spectra of Central Europe. L. Stuchi.ik, 
M. KonzalovA.— Light and scanning electron microscope study of LactucaL. and Cichorium 
L. pollen (Compositae: Lactuceae). M.Z. Haque, M.B.E. Godward. — Analisis poirnico de 

Ferrera, C.E. Hernandez Fuentes, M. Cabrera Castellanos.— Botanical identification of 

nation of their hallucinogen ergot alkaloids. L. Botz, E. Hahn, L. Gy. Szabo. 

Flora of Cuba and Antilles, IV. A. Borhidi. — A new Erythroxylum species in Cuba. R. 

species group in the genus Tabebuia Gomes ex DC. (Bignoniaceae).J. Paclt. — Ekmaniopappus 
Borhidi gen. novum (Senecioneae: Asteraceae) in Hispaniola. A. Borhidi, E. Gondar, T. 
Kiss, Zs. Orosz-KovAcs. — Is there vegetation continum in mangrove swamps? Imoh 
Ukpong.— Xylotomical study of some Venezuelan tree species (Minosaceae I-IV). K. 



Department of Botany 

University of Calicut 

Kerala 673 653, INDIA 

:omplejo Sida rombifolia en la I ndia. S. rhomboidea y 
diferentes; S. retusa, tambien restablecida como es 
s antiguo de S. alntfolta. Se ofrecen una clave art 

The genus SidaL. is taxonomically difficult. Among its species, 5. rhombifolia 
continues to be the hardest nut of all. There is great diversity of opinion as 
to its circumscription. Even as a world-wide revision of the genus is still want- 
ing, we have tended to consider this species as a highly polymorphic taxon 
including widely different elements from different areas. The result is that 
classifications proposed for one region, say for Malesia (Borssum Waalkes 
1966), Nigeria (Ugborogho 1980), Mexico (Fryxell 1988), China (Hu 
1955), and Taiwan (Chang 1977), are not applicable to Indian elements. 
This makes the current taxonomy of the species a little suspect and it seems 
that the very concept of species needs change. From the treatments of the 
complex available, it seems that defining the species in this complex into 
well characterized, easily identifiable units on narrower lines would be of 
greater service than dumping everything into a vague and hazy S. rhombifolia. 
This is the idea that emerged during a critical and exhaustive study of the 
Indian elements during a revisionary study of the Malvaceae of the area. 

Sida rhombifolia, as it is understood at present, is one of the most variable 
and widely distributed species of the genus and has been given some 30 
binomials by different taxonomists (Ugborogho 1980). Linnaeus (1753, 
1763) described three species now included in this complex, namely S. 
rhombifolia, S. alnifolta, and S. retusa, all from India. Cavanilles (1785) added 
S. alba, S. canescens, S. mkrophylla, and S. onentalis, and De CandoUe (1824) 
recognized S. capensis, S. philippica, S . plamcaulis , and S. schrankii. That was 

the time when the species had narrower definitions, but they were soon 
rejected in favor of a broad concept; consequently most such species were 
sunk into synonymy of a broad S. rhombifolta or were recognized as subspe- 
cific taxa under it. 

It was iVIasters (1874) who broadened the definition of the species; he 
recognized five different varieties of it in India. Schumann (1892) recog- 
nized six varieties in Brazil "because he had no clear idea what S. rhombifolia 
sens. str. was" (Fryxell, pers. comm.). A similar course followed by Baker 
(1892) and, for the African elements, by Hutchinson and Dalziel (1958), 
who recognized three varieties of this species in Africa. Borssum Waalkes 
(1966), working on Malesian Malvaceae, elevated var. rhombifolia and var. 
retusa into subspecies and reduced all other binomials into synonymy. This 
is the treatment widely followed at present. Subsequently, Hatusima (197 1) 
recognized another new subspecies (subsp. imularis) in the Philippines; this 
was later reported also from Taiwan (Chang 1977). Ugborogho (1980) kept 
S. retusa and S. alnifolia as two different subspecies in Nigeria. 

The classification of this complex becomes all the more difficult because 
taxonomists differ among themselves in its circumscription. For example, 
S. ortentalts Cav. has been included in S. rhombifolia by some (e.g., Rao 1985), 
but others treat it as a synonym of ^. acuta (Borssum Waalkes 1966). The 
most confusing part of all is that very different elements from different parts 
of the world have been treated under the same name by different authors. 
Thus S. alba L. is actually S. spinosa L., but S. alba Cav. is S. rhombifolia. To 
add to the trouble, Paul and Nayar (1988) treated S. alba L. and S. spmosa L. 
as two different species. So also, S. alnifolia L., widely accepted as conspecific 
with 5. retusa L., is kept distinct by Ugborogho ( 1 980), who recognized three 
subspecies in xheS. rhombifolia complex in Nigeria, namely, subsp. rhombifolia, 
subsp. alnifolia, and subsp. retusa. From his descriptions and plates, it is 
obvious that he treated at least the two latter taxa in a sense different from 
that of Linnaeus, because both of them seem to belong to subsp. rhombifolia 
(sensu Borssum Waalkes, 1966). It is pertinent to quote here the comments 
of Marais (1983) on this: "He uses some names in a sense other than that of 
van Borssum Waalkes and myself ... the photographs of the mericarps of ^. 
rhombifolia sens. lat. (p. 83 Fig. 6) are so poorly reproduced that it is 
impossible to comment on his application of the three Linnaean epithets" 
(i.e., rhombifolia, alnifolia, and retusa). 

We undertake here a revision of this complex. We have not been able to 
study it from the entire range because of difficulties in procuring materials 
and literature. Instead, we have confined ourselves to the Indian elements. 

Linnaeus (1753) described S. rhombifolia as "Sida foliis lanceolato- 
rhomboidibus serratis" and S. alnifolia as "Sida foliis orbicularis plicatis 
serratis." Later, he described another species, S. retusa (Linnaeus 1763), as 

SIVARAJAN AND Pradeep, Sida rhombifolia 65 

"Sida foliis cuneiformibus retusis, serratis subtus tomentosis. " He described 
all these taxa based on Indian specimens; while describing S. retusa, he indi- 
cated that It is very close to S. almfolia ("Varietarum S. almfoliae statuit 
Dillenms, mihi distincta visa est"). These three taxa have now been lecto- 
typified {S. rhombifolia: lectotype 2. rhombifolia LINN 866. 3, BM-Fryxell 
1 988; 5^. alnifolia: lectotype Herb. Herm. Ill fol. 4. Linn. 260. BM-Borssum 
Waalkes 1966; S. retusa: lectotype Unn. 866.7, LINN-Borssum Waaikes 
1966). We have now studied all these (as photographs) along with Indian 

S. retusa L. are conspecific. 

Sida alba, another name often reduced to the synonymy oiS. rhombifolia, 
was originally described by Linnaeus (1763) as "Sida foliis cordatis sub- 
rotundis, stipulis setaceis, axillis trispinosis" based on Indian material 
(lectotype H.U. Herb. Unn. 866.2). He also indicated its close kinship with 
S. spinosa ("Similissima S. spinosae, sed folia rotunda flores albi, stigmatibus 
purpureis; petiolarum apices purpurascentis tota demum planta major"). 
Consequently, most authors have reduced it to synonymy of S. spinosa 
(Masters 1874; Borssum Waalkes 1966; Ugborogho 1980; Fryxell 1988). 
But Paul and Nayar (1988), in their revision of Indian Malvaceae, treated 5". 
alba and S. spinosa as separate species based on presence or absence of stipular 
spines. We studied the types of S. spinosa (Lectotype: Herb. Linn. 866.1, 
LINN) and that of 5. alba L. (I.e.). They differ mainly in shape of leaves, 
probably due to morphological plasticity (as also acknowledged by Ugborogho 
1980) and in stipular spines. However, "the subpetiolar spine (better: spur) 
is quite variable in expression (even from one node to the next of a given 
plant!) and its presence or absence is of little taxonomic value. Moreover, it 
sometimes occurs on several other unrelated species, at least in South 
America" (Fryxell, pers. comm., 6 Sep 1993). Our observations on Indian 
specimens endorse Fryxell's viewpoint. These spines usually occur on the 
woody branches of older plants. The Linnaean specimen is, most likely, a 
young plant where spines have not yet developed. Otherwise, his S. spinosa 
and S. alba are identical and hence conspecific. 

CavaniUes, however, (1785) seems to have attributed the name S. alba to 
a different Indian plant (type: MA, Photo BM, not seen), which is now 
widely accepted as conspecific with S. rhombifolia. 

Sida obovata Wall, and S. microphylla Cav. from Bengal are two other names 
often associated with S. rhombifolia. In fact. Masters (1874) recognized them 
as two different varieties of the same species. We have not been able to study 
the type of the latter {Sonnerat s.n., P), but Paul and Nayar (1988) studied it, 
concluding that it is nothing but a smaller form ofS. rhombifolia. We have, 
however, been able to study the type ofS. obovata Wall. (lectotype, Wall. Cat. 
1864 from Burma, K). It is a very distinctive plant with obovate-obtuse 

leaves and long-peduncled, somev 

complex. In the absence of more material, we are not able to comment upon 
its exact identity, but are almost certain that its place is not in this complex. 

In his classification of this complex. Masters (1874) recognized five vari- 
eties of 5. rhombifolia in the Indian subcontinent, i.e., var. scabrida, var. retusa, 
var. rhomboidea, var. obovata, and var. microphylla; we have already eliminated 
the last two from the present consideration. Subsequently Borssum Waalkes 
(I966) found that Malesian materials of this complex fall under two groups, 
S. rhombifolia and S. retusa, distinguishable by habit, leaf shape, flower size, 
and some other characters, which he has dealt with in detail (Borssum 
Waalkes 1953). He also observed that, though the taxa are sympatric, they 
do not interbreed and produce hybrids in nature (Borssum Waalkes 1966). 
Consequently, he rejected Masters' treatment ("the differences between the 
groups are several, they cannot merely be regarded as varieties") and gave 
them the rank of subspecies. But his subsp. rhombifolia, still, is very highly 
polymorphic, including in it S. scabrida and 5'. rhomboidea. 

After a critical study of living populations and herbarium specimens we 
are still at a loss to understand why we cannot consider 5". retusa to be a species 
distinct from S. rhombifolia, in the strict sense of Linnaeus. The differences 
between the two. Backer (1943), Borssum Waalkes (1953, 1966), listed by 
Masters (1874), and Ridley (1922), are substantial and adequate for specific 
separation, notwithstanding the muddling by Ugborogho (1980: 65-75); 
their reproductive isolation renders them good "biological" species. We 
reinstate S. retusa as a species different from S. rhombifolia. 

But one of the problems that usually crops up in distinguishing S. retusa is 
that taxonomists often rely too much on leaf characters, especially the retuse 
leaf apex, but this character is not unique to this taxon; an unambiguous 
classification would not be possible unless we correlate it with mericarp fea- 
tures. Dr. Fryxell, while commenting on this manuscript (pers. comm.), 
wrote, "I have six specimens in my herbarium, of which three (from India and 
Ceylon) have muticous mericarps (in agreement with your description), but 
the other three have long spines (to 3 mm) on the mericarps. These specimens 
are from Malaysia and two from the Philippines. All have pubescent fruits." 
The awns of the mericarps are evident in the Malaysian specimen {Worthington 
13027 UTEP, pf), a photocopy of which Dr. Fryxell kindly sent to us. 
However, after consulting the type ofS. retusa and other Indian specimens, we 
are of the opinion that the Malaysian and the Philippine materials belong 
elsewhere than to S. retusa. But there is a nomenclatural problem involved. 

It was already mentioned that S. alnifolia and S. retusa are conspecific, the 
epithet "alnifolia" antedating the latter. There is technically nothing wrong 
with Borssum Waalkes' (1966) adoption of the later epithet "retusa" for the 

subspecies, because a name does not have priority outside its own rank 
(ICBN Art. 60), but he has also rightly suggested that "in case this taxon is 
considered a distinct species, it should be named as S. alnifolia L." So, we 
treat this taxon under the earlier name. 

Since the publication oi S. rhombtfolta by Linnaeus (1753), Fleming 
(1810) described yet another, closely related Indian species under the name 
S. rhomboidea, based on a specimen named by Roxburgh (see Borssum 
Waalkes 1966: 197; Fryxell 1988: 403). Fleming (1810) and Roxburgh 
(1832) described it as differing from S. rhombifolia "in the arils having no 
horns." But Wallich, apparently, attributed this name to a different plant 
(Wall. Cat. 1862 F) and observed that Roxburgh's plant is S. rhombifolia 
(distinguished by the two long awns on the mericarps) (cf. Wight & Arnott 
1834). It is probably from this time on that S. rhomboidea has been included 
in synonymy of the latter. However, Wight and Arnott (1834: 58) studied 
Roxburgh's figures and material cultivated in the Calcutta Garden by 
Roxburgh and concluded that they are different from Wallich 's S. rhombifolia. 
Borssum Waalkes (1966: 197) also made a critical study of these and 
concluded that "Wallich's specimen is obviously misidentified." He, in turn, 
designated the two specimens at Brussels (s. loc, s. coll., no. 2228, photographs 
seen) as the lectotype of 5^. rhomboidea. 

Sida rhomboidea is widely distributed in India, and we had ample oppor- 
tunities to study it in the field and laboratory and to compare it with S. 
rhombifolia L. (s.s), which is also plentiful. They differ substantially in several 
respects, of which fruit characters are most striking. In the case of S. 
rhombifolia, the schizocarps are prominently beaked at the apex with the pro- 
jecting awns of the mericarps. The mericarps, each with two long awns, 
dehisce at the apex, exposing part of the seed before dispersal and they 
separate from each other even as the fruits remain on the plants. In the case 
of ^. rhomboidea, however, the schizocarps are depressed, somewhat umbili- 
cated at top, as described by Roxburgh (1832), but not beaked. The 
mericarps are rather indehiscent, with a beak at apex (not 2-awned). They 
cohere together closely and are shed as a single unit by articulation of the 
pedicel, the mericarps separating much later. From our observations here, it 
is obvious that S. rhomboidea is a distinct taxon, morphologically and 
apparently reproductively isolated from S. rhombifolia and that it deserves to 
be treated as a distinct species on its own as has been done by Fleming (1810), 
Roxburgh (1832), and Wight and Arnott (1834), instead of drowning it in 
a highly polymorphic S. rhombifolia. Sida rhomboidea seems to be more akin 
to S. unicornis Marais from Mauritius. It differs in its glabrous mericarps with 
obtuse or retuse beaks, whereas S. unicornis is characterized by pubescent 
mericarps with sharp-pointed beak. 

Stda scabnda was originally described as "whole plants sprinkled with 
rigid, simple or 2-3 (or more) partite hairs . . . leaves rhomboid or oblong, 
lanceolate . . . without tomentum . . . pedicels jointed at the very base; carpels 
9-11, bicuspidate" (Wight & Arnott 1834:57), based on the peninsular 
Indian specimen Wight 1 66 (K, CAL). We have collected and studied several 
specimens that perfectly match the type (photograph seen). Apparently, it 
is close to S. rhombifolia, so much so that Masters (1 874) and Paul and Nayar 
(1988) treated it as a variety of the latter. But it differs from S. rhombifolia 
(s.s.) in a number of characters, notably in the indumentum, coarse pubes- 
cence of leaves, calyces etc. Apart from that, the mericarps in S. scabrida are 
stellate-pubescent while those of 5^. rhombifolia are glabrous. 

Sida scabrida also closely resembles S. yunnanensis Hu, with which it might 
easily be confused. But the former is characterized by adpressed simple hairs 
on the upper surface of leaves, pedicels articulated at base or not at all, and 
7-10 mericarps in contrast to the stellate hairs on the upper surface of leaves, 
pedicels articulated above the middle, and 6-7 mericarps, in S. yunnanensis. 

Now the question arises whether these are sufficient reasons for segregat- 
ing S. scabrida as a distinct species or whether we should retain it as a variety 
of 5. rhombifolia. The answer will have to focus on the manner of delimitation 
of species in the entire genus. With a circumtropical distribution and wide- 
character differences with, of course, support from apparent reproductive 
isolation. In this context, S. scabrida qualifies well for species status and so 
we are reinstating it as distinct. 

Our studies on vegetative features (from seedling stage to adult plants) 
and reproductive characters, especially mericarp morphology (Sivarajan et 
al. 1992),reveal that in India we have at least four distinct species, i.e., S. 
rhombifolia (s.s.), S. alnifolia, S. scabnda, and S. rhomboidea, which have been 
hitherto included in S. rhombifolia (s.l.). Amended descriptions and syn- 
onymy of these taxa are given below. 

c with many-rayed s 

Sida remsa L. Sp. PI. 961. 1763. Type: Herb. Linn. 866.7, LINN; Wight & Arn., Prodr. 

Fl. Pen. Ind. Or. 1:58. 1834. 
S. rhombtfolia var. retusa (L.) Mast, in Hook, f., Fl. Brit. India 1:324. 1874; Trimen, 

Handb. Fl. Ceylon 1:143. 1893. 

44. 1982; Said. & Ramesh in Said., Fl. Karnataka 1 :260. 1984; Paul & Nayar, in Nayar 
et al. (eds.X Fasc. Fl. India 19:216. 1988, non Ugborogho 1980. 
S. rhombtfolia subsp. alnifolia Ugborogho, Bol. Soc. Brot. Ser. 54:70. 1980. 
Woody herbs or subshrubs to 50 cm, usually low and strongly branched; 
stems prostate or ascending, terete, green or purplish grey, stellate-tomen- 
tose to glabrescent. Leaf blades 0.5-5 X 0.5-4 cm, leaves towards stem base 
always obovate with retuse or emarginate apex, rarely truncate, upper leaves 
obovate to elliptic-lanceolate with rounded, subobtuse or acute apex, obtuse 
or rounded at base, margins irregularly serrate-dentate or crenate distally, 
entire proximally, upper surface sparsely pubescent with short many-rayed 
stellate hairs, lower surface densely greyish tomentose with short, many- 
rayed stellate hairs; petiole 3-5 mm long, stellate-pubescent, shortly 
pulvinulate below blade; stipules 4.5 mm long, equal, linear to subulate, 
glabrescent. Flowers axillary, solitary, sometimes in terminal clusters due to 
reduction of distal leaves; pedicels longer than petioles, 3-4 mm long in 
flower, to 30 mm in fruit, glabrous, jointed about middle; calyx 6-7 mm 
diameter, 6 mm long, pubescent with minute stellate hairs, glabrous within, 
5-lobed, lobes 2 mm long, ovate-triangular; corolla 12 mm diameter, orange 
yellow, petals 7.8 x 6.5 cm, obliquely obovate, cuneate at apex, glabrous 
except for minutely hairy base; staminal column 3 mm long, glabrous or 
minutely stellate-hairy, antheriferous at apex; ovary 1.5 mm diameter, 
depressed globose, glabrous; styles 7-10, white; stigmas capitate, creamy- 
yellow. Mericarps 7-10,2.5-1.5 mm, included in calyx, reticulate or rugose 
on sides and back, apex with a pair of short stellate-hairy mucros, mucros 
obtuse, retuse, or emarginate at apex. Seeds 2 mm long, black, glabrous 
except for puberulent hilum. 

Distribution and Ecology. Sida alnifolia is widely distributed in the 
plains and hills of Southern Peninsular India, and occurs along Ghat 
roadsides, forest clearings as secondary growth, lateritic hill slopes, and 
occasionally as a weed in upland cultivation. It flowers from August to April 
in Kerala. The flowers open at about 0900 am and wither by 1400. 

FIG 1 Stdaalmfoha A Flowering branch, B-E Range ot leaf %,li uiion, 1 Indununcum; 
lateral and frontal view, K Seed, L Seedling 

Specimens examined: ANDHRA PRADESH: Cuddapah Dt.: without precise locality, 
Barher 4634 (MH). Guntur Dt.: without precise locality, Barber 4632 (MH). East 
Godavari Dt.: Bison Hill, Barber 5104 (MH). 

KARN ATAKA : Ch ikmangalur Dt. : Santaveri, PrWff/^ 4 7 732 (CALI).DakshinKaiinad 

SiVARAJAN AND PrADEEP, Sida rhombifolia 7 1 

Dt.: SuUia, Barber 2095 (MH); \]duppi,Pradeep 3060 (pf); Nalgody-Kurajadri road, Pradeep 
5041 (CALI). Kodagu Dt.: Sampaji, Barkr 2209 (MH). Uttar Kannad Dt.: Karwar, 
Pradeep 5017, 5018; Shrunkeri, Pradeep 5023B; Jogfalls, Pradeep 5041 (CALI). 

KERALA: Idukki Dt.: Munnar, Vwekanandan 48577 (MH). Kannur Dt.: Kannoth 
R.F.,Ramachandran 57700 (MH). Kottayam Dt.: Kumaramperoor R.F.,Subramaman 8246 
(FRC). Kozhikode Dt.: Kad^lundi,Pradeep5013; Badagara, Pradeep 5025 (CALI). Palakkad 
Dt.: Dhony, Pradeep 5268 (CALI); Sebastian 21092 (MH); Venkatasubramaman & Sastdharan 
7 0630 (FRC). ThrissurDt.:Peringalkuth,P?Wfe^ 5228 (CALI); Peechi,5a^r^»z^«/^K 93 22 
(FRC). Wynad Dt.: Tariodu R.R, Pradeep 51617 {ChU). 

TAMIL NADU. Coimbatore Dt.: Maruthamalai, Sebastian 721 (MH); Fischer 2017 
(FRC); Pradeep 5238, 50429 (CALI). Kottabomman Dt.: Timnelveli, Barber 535 (MH). 
Niigiri Dt.: Theppakad, Vivekanandan 4305 1 (MH). Ramanathapuram Dt.: Srivilliputhur, 
Srinivasan 72378 (MH). Salem Dt.: Hunasur cattle farm, Narayanaswami 2925 (MH). 

2. Sida rhombifolia L., Sp. PI. 684. 1753. (Fig. 2). Type: "2 rhombifolia" 
(lectotype: LINN - 866.3; isolectotype: S); Roxb., Fl. Ind. ed. Carey 3:176. 1832; 
Mast, m Hook, f., Fl. Brit. India 1:323. 1874, in part; Trimen, Handb. Fl. Ceylon 
1:143. 1893, excl. var. 3 retusa; Dunn in Gamble, Fl. Pres. Madras 1:90. 1915; 
Ramam. in Said. & Nicolson, Fl. Hassan Dr. 155. 1976; Britto & Matthew in 
Matthew, Fl. Tarn. Carnatic 3:131 . 1983, in part; Fryxell, Syst. Bot. Monogr. 25:403. 
1988, in part, excl. syn. S. rhomboidea; Mani., Fl. Silent Valley 28. 1988, in part., 

S. alba Cav., Diss. I. 22. t. 3.f.8. 1785, non Linn., 1763. 

Erect branched undershrubs to 1 m tall; stems terete, green or purplish, 
cinereous with many-armed short-stellate hairs. Leaf blades 5-6 X 2-2.5 cm, 
elliptic to rhomboid, rounded to truncate at base, 3-nerved, serrate distally, 
entire towards base, upper surface green, glabrescent, lower surface densely 
cinereous with short-stellate hairs, appearing farinaceous; petiole 5-6 mm 
long, densely pubescent, shortly pulvinulate at both ends; stipules up to 5 
mm long, equal linear-lanceolate, 1 -nerved, margins simple-hairy. Flowers 
axillary, solitary, sometimes in apparent racemes due to reduction of upper 
leaves; pedicels 0.5-1 cm long in flower, to 5 cm in fruit, glabrous, articu- 
lated above middle; calyx 5-6 mm diameter, campanulate, 10-ribbed at 
base, pubescent, 5-lobed, lobes 3 mm long, triangular, apex acute-acumi- 
nate, externally pubescent with short-stellate and simple hairs, glabrous 
within; corolla 1 cm diameter, pale yellow or creamy-white, veins some- 
times tinged with red at center; petals 9-10 X 6-7 mm, obliquely obovate, 
truncate or cuneate at apex, short stellate-hairy or glabrous at base; staminal 
column 3 mm long, glabrous, antheriferous at apex; ovary 1.5 mm diameter, 
conical, glabrous; styles 8-10; stigmas capitate, yellow or pale pink. Men- 

FIG. 2. Stda rhomhtfotta. A. Flowering branch; B-C. Rang< 

SiVARAJAN AND Pradeep, Sida rhombifolia 7 3 

carps 8-10, 4x3 mm, included in calyx, trigonous with acute angles, pale 
when mature, distinctly and transversely rugose on sides and back; apex 
with a pair of short, glabrous, divergent awns 1-1.5 mm long (as long or 
slightly shorter than calyx). Seeds 2 mm long, black, minutely hairy at hilum. 
Distribution and Ecology. Sida rhombifolia is almost world-wide in 
distribution, occurring throughout the tropics of both the Old and New 
worlds and extending into temperate zones (Fryxell 1988). It is common 
throughout India and occurs along roadsides and wastelands at altitudes 
from 1000 to 2000 m. It flowers almost throughout the year, the flowers 
opening late in the morning, usually between 1045 and 1100. 

Specimens examined. KERALA: Kottayam Dt.: Sabarigiri, Sivarajan 5211 (CALI). 
Thiruvananthapuram Dt.: Ponmudi, Pradeep & Sivarajan 30463 (CALI); Bonecaud, 
Mobanan 34732 (MH). 

TAMIL NADU: Kanyakumari Dt.: Kothayar, Pradeep 44923, 44916 (CALI). 
Kattabomman Dt.: Courtallum, Pradeep 3103 (CALI). 

3. SidarhomboideaRoxb. exFlemmg,Asiat. Res. 11:178. 1810. (Fig. 3). 

Type: s. loc, s. coll. 2228 (BR); Roxb., Hort. Beng. 50. 1814 & Fl. Ind. ed. Carey 

3:176. 1832; Wight & Arn., Prodr. Fl. Pen. Ind. Or. 57. 1834; Dunn m Gamble, FL 

Pres. Madras 1:90. 1915. 

Sida rhombifolia var. rhomhoidea (Roxb. ex Flemmg) Mast, in Hook, f., Fi. Brit. India 

.S~. rhombifolia subsp. rhombifolia sensu Borss., Blumea 14:193. 1966; Mani. & Sivar., Fl. 
Calicut 44. 1982; Said. & Ramesh in Said., Fl. Karnataka 1:259. 1984; Nair & Nayar, 
Fl. Courtallum 1:75. 1986; Fryxell, Syst. Bot. Monogr. 25:403. 1988; Mani., Fl. Silent 
Valley 28. 1988; Paul & Nayar in Nayar et al. (eds.), Fasc. Fl. India 19:214. 1988; 
Vajravelu, Fl. Palghat Dt. 84. 1990 (all in part). 
Erect much branched subshrubs to 2.5 m tall; stems terete, usually pur- 
plish, minutely pubescent with small stellate hairs. Leaf blades on younger 
shoots much larger (7-8 X 5-6 cm), obovate or suborbicular, truncate or 
rounded at base, subobtuse or acute at apex; those on flowering shoots 
smaller, 1-5 X 0.5-3 cm, rhomboid to lanceolate, 3-nerved from base, lateral 
nerves 3-5 pairs, nerves raised on lower surface, margins coarsely serrate to 
crenate, entire towards base, densely stellate-tomentose beneath, sparsely 
pubescent above; petiole 1-15 mm long, stellate-pubescent, shortly 
pulvinulate at both ends; stipules 9 mm long, equal, linear, slightly pur- 
plish, caducous. Flowers axillary, solitary; pedicels 6 mm in flower, to 30- 
40 mm in fruit, filiform, glabrous, articulated at about middle; calyx 9 mm 
diameter, campanulate, 10-ribbed at base, 5-lobed, lobes 3 mm long, tri- 
angular, outer surface densely tomentose with minute stellate hairs, inside 
nearly glabrous, margins purplish; corolla 1 . 5 cm diameter, pale yellow; petals 
7-8 X 5-6 mm, obliquely obovate, retuse or emarginate at apex, glabrous; 
staminal column 3 mm long, stellate-pubescent, antheriferous at apex; 

ovary 1.5 mm diameter, depressed globose, glabrous; styles 8-10; stigmas 
capitate, yellow. Mericarps 8-10, completely included in calyx, closely co- 
herent, 3x2 mm, indehiscent, trigonous with acute angles, prominently 
reticulate on sides, reticulate or rugose on back, apex beaked with a single, 
glabrous muticous process. Seeds 2 mm long, brownish black, glabrous 

Distribution and Ecology. Sida rhomboidea is common in the plains of 
Peninsular India. It occurs along roadsides and waste lands generally at sea 
level. Because of its frequent confusion with S. rhombifolia, it is difficult to 
draw conclusion on its distribution elsewhere, based on literature. The plant 
flowers almost throughout the year. The flowers open between 1045 and 
1100 and wither by 1415. 

Specimens examined. KERALA: Ernakulam Dt.: Wellington Island, Pradeep 3203; 
Cochin, Pradeep 3179 (CALI). Kannur Dt.: Tellichetty, Punnol, Pradeep 3107 (CALI). 
Kottayam Dt.: Changanacherry, Antony 904 (MH). Kozhikode Dt.: West Hill, Pradeep 
321 3. 6039; Dtyiigin J ay a kumar 884; Chemvannur, Sivarajan 1482 (CALI). Malappuram 
Dt.: Parappanangadi, Pradeep & Sivarajan 5002 (CALI). Thiruvananthapuram Dt.: 
Bonecaud, Mohanan 63246 (MH). 

TAMIL NADU: Ramanathapuram Dt.: Vajravelu 33833 (MH). 

4. Sida scabrida Wight & Arn., Prodr. Fl. Pen. Ind. Or. 57. 18.34. (Fig. 4). 
Type: Peninsular India, Wight 166 (CAL, K,); Ugborogho, Bol, Soc. Brot., 54:100. 

Sida rhombifolia var. scabrida (Wight & Am.) Mast, in Hook, f., Fl. Brit. India 1:324. 

S. rhombifolia subsp. rhombifolia var. scabrida (Wight & Arn.) Mast., Paul & Nayar in Nayar 

et al. (eds.), Fasc. Fl. India 19:216. 1988. 
Erect branched subshrubs to 2 m tall; stems terete, green or slightly tinged 
with purple, pubescent with minute stellate hairs intermingled with 
scattered, long, simple hairs. Leaf blades 6-8 X 3-4 cm, concolorous, rhom- 
boid or oblong-lanceolate, truncate at base, acuminate at apex, basally 3- 
nerved, lateral nerves 4-5 pairs, margins serrate-crenate distally, entire 
towards base, coarsely pubescent on upper surface by appressed simple hairs, 
on lower surface with few scattered 2-3 armed stellate and simple hairs 
especially on the nerves, never with tomentum; petiole 0.5-1 cm long, 
pubescent with minute stellate and long simple hairs; stipules 4—6 mm 
long, subulate or filiform, ciliate. Flowers axillary, solitary, sometimes in 
clusters of 3— 5 due to reduction of distal leaves. Pedicel up to 1 cm in flower, 
to 3 cm in fruit, not articulate; calyx 6-7 cm diameter, 10-ribbed at base, 
campanulate, 5-lobed, lobes 4x4 mm, ovate, acuminate at apex; corolla 1 . 5 
cm diameter, yellow; petals 1-12 X 7-8 mm, minutely stellate-hairy at base; 
staminal column up to 3 mm long, stellate-pubescent; ovary ovoid, 2 mm 

ew, K Seed, L Seedl 

diameter, minutely simple-hairy towards apex; styles 7-10; stigmas sub- 
globose. Mericarps 7-10, 3 x 3 mm, enclosed in calyx, trigonous with acute 
angles, black when mature, minutely stellate-pubescent on back towards 
apex, prominently reticulate or transversely rugose on sides and back, apex 
with a pair of linear divergent simple and stellate-pubescent awns. Seeds 2 
mm long, blackish or brown, pubescent at hilum. 

Distribution diwd'EcoiogY -Sidascabrida is known from Peninsular India 

SiVARAJAN AND Pradeep, Sida rhombifolia 77 

and (disjunccly) from Nigeria in Africa. It occurs in disturbed sites in 
evergreen, semi-evergreen, and moist deciduous forests. It also grows along 
roadsides generally at an altitude of 250-2000 m. The plant produces flowers 
principally from August-April. The flowers open between 1045 and 1 100. 

Specimens examined. KARNATAKA:. Chikmagalur Dt.: Bababudan Hills, Pradeep & 
Sivarajan 47726 (CALI). Kodagu Dt.: Mercara, Sampaji, Barber 2313 (MH) 

KERALA: Kottayam Dt.: V^im^?^, Subramanian 9969 (FRC). Kozhikode Dt.: Badagara, 
Pradeep 3101 (CALI, pO. Malappuram Dt.: Nedunkayam, Pradeep & Sivarajan 3097 
(CALI). Palakkad Dt.: Silent Valley R.F., Nair 64374, Vajravelu 26103 (MH). 
Pathanamthitta Dt.: Moozhiyar, Subramaman 9620; Mahadevan & party 9193 (FRC); Anil 
Kumar 1313 (MH). Thiruvananthapuram Dt.: Ponmudi Hills, Mohanan 32342 (MH). 
Thrissur Dt.: Peringalkuthu, Pradeep 3227 (CALI). Wynad Dt.: Lakidy, Pradeep 3137, 
3134; near Pookkottu lake, Pradeep 30437 (CALI); Chandanathode, Ellis 29438 (MH). 

TAMIL NADU: Coimbatore Dt.: Attakkati-Valparai road, Sivarajan Pradeep 47761 
(CALI). Kanyakumari Dt.: Kothayar, Pradeep 44929 (CALI). Nilgiri Dt.: Devala R.R, 
Vajravelu 42831; Kattaicombai, Subramanyam 1080; without precise locality, Wigk s.n. 
(MH). Salem Dt.: Salem, Deb 31263 (MH). 


We are indebted to Dr. Paul A. Fryxell, Texas A&M University, for com- 
ments on an earlier version of our paper and for useful suggestions; to Dr. P. 
Bamps, National Botanical Garden of Belgium, for some of the type photo- 
graphs; to Dr. R.E. Ugborogho, University of Lagos, Nigeria, for help with 
literature; and to the Jt. Director, Botanical Survey of India, Coimbatore, for 
providing facilities to study materials at MH. AKP is grateful to the Council 
of Scientific and Industrial Research, New Delhi for a fellowship. 

Baki;r, E.G. 1892. Synopsis of genera and species ofMalveae. J. Bot. 30:1 36-142, 235-240, 

290-296, 324-332. 
BoRSSUM Waalkes, J. van. 1953. Enkele bloeiwaarnemingen bij twee Sida-soonen. Trop. 

Natuur 33: 21-29. 

1966. Malesian Malvaceae revised. Blumea 14:1-213. 

Candolle, A.R de. 1824. Malvaceae. In: A.R de Candolle (ed.), Prodromus 1:427-474. 
Cavanilles. A.J. 1785-1790. Monodelphiae classis dissertationes decern. Pans & Madrid. 
Chang, Ching-En. 1977. Malvaceae. In: Hui-LinLietal. (eds.) Flora ofTaiwan 3:7 10-728. 

Hindustani and Sanskrit languages. Asiatic Res. 11:153-196. 
Fryxell, RA. 1988. Malvaceae of Mexico. Syst. Bot. Monogr. 25:1-522. 
Hatusima, S. 1971. Flora of the Ryukyus. Naha, Okinawa. 
Hu, S.Y. 1955. Flora of China, family 153: Malvaceae. Jamaica Plain: Arnold Arboretum 

of Harvard University 
Hutchinson, J. and J.M. Dalziel. 1958. Flora of West Tropical Africa. Ed. 2. London, 

Vol. 1(2). 
Linnaeus, C. 1753. Species plantarum. Stockholm. 

Marais, W. 1983. Notes on Mascarene Malvaceae. Kew Bull. 38:41-46. 

Masters, M.T. 1874. Malvaceae. In: J.D. Hooker (ed.), Flora of British India 1:317-35: 

Paul, T.K. and M.P. Nayar. 1988. Malvaceae. In: M.P. Nayar, K. Thothathri, and Iv 

Sanjappa (eds.), Fascicles of flora of India. 19:64-233. 
Rao, R.S. 1985. Malvaceae. In: Flora of Goa, Dm, Daman, Dadraand Nagarhaveli. 1:31-4? 
Ridley, H.N. 1922. The flora of the Malay Peninsula 1:253-260. 

Schumann, K. 1892. Malvaceae I. In: Urban, I. (ed.) Flora Brasiliensis 12(3):253-456. 
Sivarajan, V.V., A.K. Pradeep and E.J, George. 1992. Mericarp morphology of India 

species ofSida L. (Malvaceae) m relation to taxonomy. J. Taiwan Mus. 45:65-73. 
Ugborogho, R.E. 1980. The taxonomy ofSida L. (Malvaceae) in Nigeria. I & II. Bol. Soc 

Brot., Ser. 2, 54:5-40, 65-85. 
Wight, R. and G.A.W. Arnott. 1834. Prodromus florae peninsulae Indiae orientali; 



Department of Biological Sciences 

Saint John's University 
Jamaica, NY 11439, U.S.A. 


Plant Sciences Doctoral Subprogram 

Lehman College, City University of New York 

Bronx, NY 10468, U.S.A. 

We report a scacistical analysis of leaf form for a sample of 48 1 collected from five yaupon 
trees at Ponchatoula, Louisiana. Considerable variation exists within these "sun" leaves. 
Across the 5 trees, surface area varies 24-fold (22.1-533 mm^), leaf specific mass 22-fold 
(7.67-167.83 g m^), leaf length 5-fold (6-31 mm), leaf width 4.5-fold (4-18 mm), and 
crenations 4.7-fold (7-33 per leaf)- Leaf complexity (LC) varies from 1 to 6 Fourier 
frequencies, and the leaf dissection index (DI) varies from a nearly circular 1.036 to a high 
of 1 .349. Trees in this population are statistically significantly different from each other in 
average leaf size, leaf mass, leaf specific mass, and in number of crenations per leaf. In logistic 
regression, the probability that a leaf will develop more crenations, increases with leaf size. 
As leaf size changes, shape remains relatively constant within this population; amount of leaf 
dissection does not correlate with other morphological variables. Leaf surface area is strongly 
related to nodal position on the shoot, middle nodes generally produce the largest leaves. 
Key Words: Ilex vomitona, yaupon, leaf form 


Se realiza un analisis estadistico de la forma de la hoja sobre una muestra de 481 hojas 
colectadas en cinco arboles en Ponchatoula, Louisiana. Existe una variacion considerable 
entre esta hojas de "sol." En cinco arboles, el area de la superficie varia en 24 veces (de 22.1- 
533 mm^), la masaespecfficade la hojaen22veces(de7.67-l67.83gm-2),lalongitud foliar 

crenaciones en 4.7 veces (de 7-33 por hoja). La complejidad de la hoja (LC) varia de 1 a 6 
frecuencias de Fourier, y el I'ndice de diseccion foliar (DI) varia desde casi circular 1.036 hasta 
1 .349. Los arboles de esta poblacion muestran diferencias significativas estadisticamente en 
el tamano foliar medio, masa foliar, la masa foliar especifica, y en el niimero de crenaciones 
por hoja. En regresion logistica, la probabilidad de que una hoja desarrolle mas crenaciones 
aumenta con el taman de la hoja. Cuando varia el tamano de la hoja, su forma permanece 
relativamente constante en esta poblacion; la diseccion de la hoja no se correlaciona con otras 

posicion nodal en la rama, los nudos medios producen generalmente las hojas mas grandes. 

Advances in computer-assisted image analysis expand the ability of bota- 
nists to use large sample sizes in leaf morphometric research (e.g., Kincaid 
and Schneider 1983, White et al. 1988). Our objective was to quantify leaf 
variability in a population of yaupon, Ilex vomitoria Alton (Aquifoliaceae) at 
Ponchatoula, Louisiana. We collected leaves from five trees in order to 
answer these questions: (1) Are trees homogeneous in leaf size and shape? (2) 
Is leaf form related to nodal position along the twig? (3) Do predictive 
relationships exist among leaf specific mass (g dry weight / m^ surface area), 
crenation number, mass, area, dissection index, and leaf shape complexity? 
(4) How do average leaf images per tree, reconstructed by Fourier transform, 
compare to conventional morphometric statistical analysis? 

Yaupon is a shrub and small tree common in forests along the Coastal 
Plain from southern Virginia to Florida, and west to Texas (Elias 1980). The 
leaves are small, flat, coriaceous, evergreen, elliptical, and have marginal 
mucronate crenations (Radford et al. 1968). 

A sample of 481 leaves was collected, on August 12, 1989, from 5 trees 
growing within 100 meters of each other along a sunlit edge of a pine forest 
at Ponchatoula, Louisiana. Leaves were individually numbered with a serial 

and the blades placed into a plant press. After drying in a convection oven 
at 70°C, the leaves were weighed to the nearest 0.0001 gram. Crenations 
were counted using a stereo dissecting microscope. Maximum length and 
width were recorded for each blade. 

In our laboratory, leaf images are analyzed (Fig. 1) using the leaf boundary 
method of Kincaid and Schneider (1983) which is based on Fourier transform. 
In an analysis of various computerized leaf morphometric methods, White, 
et al. (1988) found this method performed well in terms of discriminating 
power and in the reconstruction of synthetic, average leaf images. 

Leaves were photographically enlarged (Fig. 2), and the images bound- 
aries digitized into x, y coordinates using a graphics pad (Model CR1212, 
Summagraphics Corp., Fairfield, CT) attached to a Macintosh Ilci computer 
(Fig. 1). Other details are in Kincaid and Schneider (1983), and in Figures 
1-3. Image information lies in the values of the Fourier coefficients at each 
frequency (Table 1). Leaves have the same size and shape, if and only if, their 
Fourier coefficients are identical (Kincaid and Schneider 1983). Using this 
method, leaf surface area, leaf complexity, and leaf dissection index were 
computed for 342 out of the 48 1 leaves. Leaf complexity (LC) is a dimension- 
less and discrete, ordinal variable providing a mathematical measure of the 
"complexity" of a leaf's outline. For example, LC = 1 for pure ellipses, and 

The Leaf Boundary Method 

Pressed Leaf or 
Photocopy Placed 
onto Digitizer Pad 




1 performed on these p 
s that completely 
the leaf image. These coefficients are sto 
disk in a file for each leaf. 


equally spaced points (e.g. 256, 
512, 1024) along the boundary. 

frequencies (M) Reconstructioi 


M = 3reconstructsleaf#30to 

within 9 


between M= 16 and M = 32, a 

marginal detail. For leaves of SI 

mple shai 

image analysis of mucronate 

LC = 1 for most elliptically shaped leaves with relatively smooth margins. 
The more complicated the leaf shape, relative to the best fit ellipse, the 
greater the integer value of LC (In terms of reverse Fourier transform, LC is 
the number of frequencies necessary to reconstruct the leaf image to within 
95% of the actual image.). 

While LC captures a mathematical aspect of leaf shape, dissection index 
(DI) is the empirical relationship between leaf perimeter and leaf surface area 

amenable to biophysical interpretation of convective heat exchange. The 
minimum value of DI is that of a circle for which DI always equals 1 .0. The 
value of DI for a circularly shaped, entire leaf is slightly larger than 1 .0, and 
the more deeply lobed, dissected or lanceolate a leaf shape, the larger the 
value [DI = Perimeter / {2*SRQ(Area * pi))}. These variables, defined by 
Kincaid and Schneider (1983), are useful in making comparisons of leaf 
shape among leaves having different surface areas. 

We used Stat View (Abacus) and JMP (SAS 1 989) on Macintosh comput 
ers to perform the data analysis. The technique of logistic regression (Pagan( 
and Gauvreau 1993; using JMP) was utilized to search for trends, and u 
visualize relationships between crenation interval (dependent variable) anc 
leaf surface area, and between classification of leaves by tree (dependen 
variable) and number of leaf crenations. While logistic regression is ; 
commonly used statistical tool in biomedical fields for analyzing discreti 
responses, it is rarely used in organismic biology. 

mm2), leaf specific mass 22-fold (7.67-167.83 g m'^), leaf length 5 -fold (6- 
3 1 mm), leaf width 4.5-fold (4-18 mm), crenations 4.7-fold (7-33 per leaf), 
and leaf complexity (LC) from 1 to 6 Fourier frequencies. Leaf dissection 
index (DI) changes from a nearly circular 1.036 to a high of 1 .349- Table 2 
provides descriptive statistics and results of analysis of variance for these 
leaves grouped by tree. 

We found strong evidence for heterogeneity among the five trees for 
average leaf area, mass, leaf specific mass, crenation number, and dissection 
index (P < 0.00001 with R-square values for the main effect ranging from 
0. 153-0.542, Table 2). Interestingly, average LC was homogeneous among 
the 5 trees (F = 1.7; 4,338 df; P = 0.l4) and homogeneous among shoots 
within each tree (P > 0.05). We predicted that leaf dissection, a variable 
providing linkage to convective cooling ability, would be positively corre- 
lated with leaf surface area. However leaf dissection did not correlate with 
any variable, indicating that as leaf size changes, shape remains relatively 
constant within this population. 

In these "sun" leaves, leaf weight (r = 0.91), leaf specific mass (r = 0.24), 
and crenation number (r = 0.48) increased with leaf size (P < 0.01 for each 
correlation coefficient). Figure 4 presents the relationship between leaf mass 
and leaf area. On an individual tree basis, dry weight of leaf is an excellent 
predictor of surface area (e.g., in Figure 5 for Tree 5, R-square - 0.976) but 
less so for all leaves (R-square = 0.826). As a field technique, leaf width pro- 
vides the simplest predictor of leaf surface area (e.g., for Trees 1 & 4, Area = 
25.30 * Width - 60.46, R-sq. = 0.91). Once it is determined that leaf shape 
changes little with leaf area, regression equations could be used to predict 
leaf area, as dry weight and/or blade width is easier to measure than leaf area. 

Increased leaf specific mass usually confers greater water use efficiency and 
photosynthetic capacity. Average leaf specific mass ranged from 82.6 in tree 
4, to 141.3 g m"2 in tree 5. We cannot explain why leaf specific mass varied 
so much in this study, among 5 trees growing within 100 meters of each 
other (Table 1). Indeed, trees 4 and 5, with essentially the same average leaf 
images (Fig. 8), had the most divergent values for leaf specific mass. 

Crenation number, grouped into 5 levels (7-10, 11-14, 15-18, 19-22, 
and 23-33 crenations per blade), was declared a "response" variable, and 
analyzed by logistic regression against leaf surface area as an explanatory 
variable (Figure 5). In logistic regression, crenation value is not predicted, 
rather, probabilities are estimated for each level of crenation "response," 
given leaf surface area. The resultant graph, partitions the outcome space 
into mutually exclusive regions. To use the logistic regression graph, one 
draws a vertical line at any desired surface area dividing the estimated 
probability into segments for each level of response. For example, as leaf size 
increases beyond 200 mm^ the probability that a leaf will have 7-10 

J5 lAu- 
'% WO. 

J.424X + 17.956, R-squared: .976 

Mass (grams dry weight) 

ikely . 

-33 c 

iikely to develop " 

In Figure 6, we use logistic regression as a tool to visualize the estimated 
probability of tree "membership" for a leaf, given the number of crenations 
on a particular leaf Trees 1 and 3 have the lowest average number of 
crenations per leaf; and Trees 2 and 4 have the largest average number (Table 
2). The fitted logistic regression curves of Figure 6 provide a display of these 

Leaf surfac 



, is strongly related to nodal : 

but only wh 



\ \ 23-33 


^ 0.75- 





\ 19-22 \ 

\ crenations \ 

-C5 0.50- 






\ crenations \ \ 



\ \ \ 

LU 0.25- 

\ ere 

nationsX \ \ 


7-10 \ 

^^ \^ 




Leaf Surface Area (sq. mm) 

analyzed on a per shoot basis (Fig. 7). For many siioots, 70-90% (R-square) 
of total variability in leaf area is accounted for by nodal position. This holds 
true even for long terminal shoots that have experienced, over the grov^^ing 
season, an episodic growth and/or a developmental switch from preformed 
to neo-formed (produced and released in current growing season) leaf buds 
(lower right graph in Fig. 7). Shoots within trees had the same average leaf 
size in ANOVA. 

Average leaf images reconstructed for each tree, as computed from average 
Fourier coefficients for the first 8 frequencies, are displayed in Figure 8 along 
with principal component analysis of more conventional morphological 
variables. The Fourier transform captures only the two-dimensional leaf 
outline: We see from these average images that Trees 4 and 5 have very 
similar leaf sizes and shapes, and that Trees 1 and 3 have similar leaf sizes but 
different shapes. In principal component analysis, leaf area loaded heavily on 
the first component which accounted for 0.33 of total variance. The shape 
variables (DI and LC) loaded on the second component, accounting for 0.28 
of the variance. Tree number, loaded on the third component, accounting for 
0.19 of the variance. As a general rule for biological objects analyzed by 






= 0.75- 



Tree 4 > 






T3 0.50- 


\ ^/'^ 






^^:::^— -s^ 

Tree 2 

LU 0.25- 

Tree 1 


^ -__ 

5 10 15 20 25 30 35 

Number of Crenations on Leaf 

:ipal componej 

.riables domi 

priutipai coiiipuueiics, size variaoies auiiiiiiace tiie iirst compone 
"shape" variables the second compo "' ' ~" 

the 342 leaves, graphed in PC spac 

11979). The < 

bivariate means or each or t\ 
Figure 8. Trees 3, 4 and 5 cl 

, graphed in PC space for the first two components, revea 
spending to tree. The 95% confidence ellipi 
■ ofth ' 

are displayed 

rather closely toget 

, while Trees 1 and 

Leaf surfiice area appears to be functionally related to nodal posi 


; at the early and late nodes (Fig. 7). This range in leaf 
igle shoots, may transcend average differences from 
latitudinal and sun-shade extremes of the species. 

FIG. 7. Leaf surf 
rcent (R-square i 
:ounted for by no< 

ower right figure 

We have established nine study populations of yaupon. Three of these are 
inland stations: Aiken, South Carolina; Homosassa Springs, Florida; and, 
the subject of this paper, Ponchatula, Louisiana. We have also established six 
coastal stations from the species northern distributional limit, Virginia 
Beach, Virginia, to one of its southern limits. New Smyrna Beach, Florida. 
Exploratory analysis of the modestly sized data set of this paper has helped 
us plan collection strategies for the other stations. As botanical methods of 
computer-assisted image analysis advance (White et al. 1988), research 
designs should be able to accommodate larger sample sizes of leaves, 
involving more trees per population, and more shoots per tree. 

For a grant in support of field work, we thank the Research and Develop- 
ment Committee of the Holly Society of America, Inc., Barton Bauers, Sr., 
Chairman. We also thank Maria Bilelis, iVfark Pokrzywa, Josephine Puppio 
and Robert Sica, students at St. John's University, Jamaica, New York for 
morphometric assistance. 

Tree 1 Tree 2 Tree 3 Tree 4 Tree 5 

Small are 


Large are 



Tree 4 


Tree 2 

Tree sV 


Vy Tree 3 



FIG. 8. Upper: Average leaf images 
coefficients. Lower: Principal component 

from average Fouri 

Elias, T.S. 1 980. The complete trees of North America. Van Nostrand Reinhold, New York. 
KiNCAiD,D.T. and R.B.Schneider. 1983. Quantification of leaf shape with amicrocomputer 

and Fourier transform. Canad. J. Bot. 61:2333-2342. 
Pagano, M. and K. Gauvreau. 1993- Principles of biostatistics. Duxbury Press, Belmont, 

PiMENTEL, R.A. 1 979- Morphometries the multivariate analysis of biological data. Kendall/ 

Hunt Publishing Co., Dubuque, Iowa. 
Radford, A.E., H.E. and C.R. Bell. 1968. Manual of the vascular flora of the 

Carolinas. University North Carolina Press, Chapel Hill. 
SAS. 1989. JMP User's Guide. SAS Institute Inc., Cary, North Carolina. 
White, R.J., H.C. Prentice and T. Verwijst 1988. Automated image acquisition and 

morphometric description. Canad. J. Bot. 66:450-459- 



Department of Botany 

Duke University 

Durham, NC 27708, U.S.A. 

■ compelling case for recognizing three genera of North American Myricacea 
'oma, More/ia, and Myrka) is presented. Keys and descriptions are provided for th 
lized supraspecific taxa. The nomenclatural basis of each of these genera, subgenera 

nd in the Americas and Africa and subg. Morella is restricted to eastern an( 
■astern Asia. There are at least four species oi Morella in the United States and Canad 
■rhaps as many as six: M. cenfera (L.) Small, M. caroltmemts (Mill.) Small, M. inodon 
irtram) Small, and M. caltformca (Cham. & Schltdl.) Wilbur. Those about which ther 
le question do not have binomials in the genus Morella but are known as Myrid 
'anica Mirbel (= Cerothamnus pensylvamcus (Mirbel) Moldenke) and Myrkapusilla Rai 
ithamnus pusilla (Raf.) Small). Myrica L. is represented by the circumboreal M. gale L 

•grina (L.) J.M. Coulter, is restricted largely to the northeastern United States an< 

a subg. Cerothamnus (Tidestr.) Wilbur and series Faya (Webb & Berthel.) Wilbui 
a califormca (Cham. & Schltdl.) Wilbur, and for the Azorean M. faya (Alton) Wilbui 



udio convir 


tres generos de Myricaceae de Norte 

Morella, y 

claves y desc 

ripciones de los taxa 

supraspecificos reconocidos. Se bosqueja y discute 1 

a base nomen 

clatural de todos los 

generos, subgeneros > 

el genero mas amplio, 

ibgeneros de los cuales Cerothamnm 

■ (Tidestr.) Wilbur se 

encuentra tanto en . 

America d( 

,\ Norte como del Sur y en Afria 

I, y Morella que esta 


ureste de A 

sia. Hay al menos cuat 

:ro especies de 

Morella en los Estados 

Unidosy Canada, que 

quizas puedan llegar a seis: M. ceri 

■/"era (L.) Small, 

M. caroliniemis (Mill.) 

Small, M. inodora {\ 

V. Bartam) 

Small, y M. californi 

■ca (Cham. & : 

Schltdl.) Wilbur. Las 

especies sobre las que 

= hay dudas 


nomen en el genero Morella, pero se 


■a Mirbel (= Cerotham^ 


<s (Mirbel) Moldenke) 


= Cerofham; 




iW.L., circumboreal, 


.^i/S. Watson, deCalii 

fornia. Compton 

su unica especie C. p 

^eregnna (L. 

)J.M. Coulter estar 

loreste de los Estados 

Unidos y zonas adya 

:anada. Se dan nuevaj 

i combinacion( 

;s y/o ranges para los 

'ilbur y para M. faya ( 

94 S!DA16(1) 1994 

For one whose formative years were spent in a section of the country where 
Small's ( \9^y) Manual of the southeastern Flora was the basic reference, numer- 
ous adjustments had to be made to relate to the prevailing, more conservative 
generic concept then dominating most of American botany. Small's "micro- 
genera" were viewed as a provincial aberration. For one's work to be under- 
stood by most of the botanical community, one had to convert the generic 
names employed by Small and other prolific practitioners of the so-called 
"New York School" such as Britton, Rydberg, and Barnhart into the broader 
generic concepts employed by more conservative botanists. For example, how 
many of us know which common genera are intended when one encounters 
such generic names as Wallia, Cerothamnus, Tulipastrum, or P henianthus'^ 
Brandegee (1901) argued vehemently against the generic splitting charac- 
teristic of the New York Botanical Garden under the leadership of Nathaniel 
Lord Britton. She suggested that genera should be so broadly delineated that 
all reasonably bright 10 year olds could be expected to know the principal 
genera of plants and animals in their neighborhood. Clearly Brandegee 
would be disappointed today for not only would most school boys and girls 
fail her test but so would most university biology professors. In fact, in the 
past 2 decades the pendulum has swung back strongly towards the generic 
standards of Britton, Rydberg, and Small. Who could have predicted 2 or 
3 decades ago that the well-known genus Cassia would disappear from the 
flora of the Carol inas and be replaced by the segregates Chamaecrista and 
Senna; or that Psoralea would be confined to southern Africa and that those 
generic names that Rydberg and Small were castigated for using instead 
{Orbexilum, Pediomelum, and Rhytidomene) would now be very widely adopted 
for different elements formerly included in Psoralea s.l.? 

Turning to Myricaceae, we find that Small (1 93 .3) recognized three genera 
in the area of the southeastern United States: Myrka L. (1753), Comptonia 
L'Her. ex Alton (1789) and Cerothamnus Tidestr. (1910). Radford et al. 
(1968) also recognized three myricaceous genera in the Carolinas, differing 
however in the generic names accepted and the groups to which the names 
applied: Gale Adans. (1763), Comptonia L'Her. ex Alton (1789), and Myrka 
L. (1753). More conservative treatments of Myricaceae recognize, in addi- 
tion to the controversial and relatively little-known, monotypic. New 
Caledonian Canacomyrka Guillaumin, two genera sometimes combined as 
subgenera or sections within the broad concept of Myrka. Under such a 
conservative scheme, the genus Myrka would consist of three major taxa of 
very unequal size. The New Caledonian Canacomyrica will not be considered 
in this paper, but it should be noted that considerable doubt has been 
expressed as to its relationship with Myricaceae (Thorne 1973). The 
nomenclature of the three North American myricaceous genera is in part 
controversial as is discussed below. 

The morphological evidence summarized in Table 1 argues strongly that 
there are at least three major groups within non New Caledonian Myricaceae. 
These three groupings are so fundamentally different that, in my opinion, 
generic rank should be accorded to each of them. With flowers as greatly 
reduced as those of Myricaceae, it is not surprising that the most striking 
differences are found elsewhere — especially in characteristics of fruit and 
vegetative features. These same groups were treated as three subgenera of 
Myrica by Engler (1894). Gleason (1952) and Gleason & Cronquist (1963) 
treated all species as Myrica with no indication given of infrageneric, supra- 
specific classification. Many authors including Rehder ( 1 949), Fernald (1950), 
Hutchinson (1964-), Elias (1971), and Gleason & Cronquist (1991) in the 
past 5 decades have recognized two genera: Comptonia and Myrica (sometimes 
the latter with the two commonly accepted subgenera, Gale and Morella). 
Among previous authors recognizing three genera are Chevalier (1901), 
Rendle (1903), Small (1933), Radford et al. (1968), and Baird (1968). 

That there are three major groupings within Myricaceae seems generally 
agreed upon by most students of the family; the only question is the rank or 
ranks to be accorded to these seemingly natural, monophyletic taxa. Perhaps 
a comparable case is Quercus, since that genus is traditionally divided into 
three subgenera by most botanists although Schwarz (1936) treated the 
subgenera as genera and Oersted (1867) had originally treated those Asian 
species with connate cupular scales forming concentric rings as the genus 
Cyclobalanopsis. A table or chart comparing the differences between these 
three fagaceous taxa is both lengthy and impressive, but evaluation of the 
differences suggests that they are variations on the same theme as opposed 
to rather dramatic innovations and new themes expressed in the phyletic 
development of the three lines of Myricaceae. 

Although in Table 2 Baird has been outvoted 3 to 1, the preponderance 
of evidence suggests that he was nomenclaturally correct in the names he 
employed and the groups to which he applied the names. Small (1903) 
recognized both Comptonia and Morella in the southeastern United States but 
was unaware of the presence of Myrica gale there. It is unfortunate that 
Baird's detailed investigation was never published. I here acknowledge my 
indebtedness to Baird (1968) for information summarized in his unpub- 
lished dissertation that strengthened the case made for the recognition of 
three genera. It must be admitted that in such matters one can not be 
dogmatic for the state of our science does not totally exclude individual taste. 
As long as the groups recognized are seemingly monophyletic (in the pre- 
cladistical sense) and perhaps reasonably equivalent in morphological 
differentiation, whether such groups are treated as three genera or three 
sections or subgenera of one genus is a matter of individual taste doubtlessly 
tempered by both tradition and the prevailing philosophy of the period. 


In recognizing genera, botanists would perhaps find that by placing 
greater emphasis on the currently minimized morphological adaptations 
between plants and their environment, the disparity between botanical and 
zoological practice would be less striking than now is the case. The impor- 
tance of the genus as an indicator of discrete evolutionary lines would be 
enhanced if the obvious correlation between functional morphology and 
broadly conceived environmental integration were emphasized (Inger 1958). 
In the case of Myricaceae, it should be pointed out that these morphological 
adaptations to differing means of fruit dispersal have not been ignored by 
botanists even if they have not emphasized the functional role. These charac- 
teristics of the fruit have been recognized by even the most conservative 
botanists in their formal classification but usually at the subgeneric level. 
My account merely advocates recognition at the generic level. 

A principal reason for the widely divergent generic treatments of Myricaceae 
is not because of differences in interpretations of biological or morphological 
facts or even in the weight given to these facts; it is simply a disagreement as 
to the lectotypification of the generic name Myrica L. Chevalier (1901), Rendle 
(1903), and Radford et al. (1968) in effect all treated Myrica cerifera L. as the 
lectotype of the generic name Myrica; Baird (1968) accepted Myrica gale L. as 
the lectotype. This confusion exists whether we accept one genus with three 
equivalently ranked subgroups or recognize three independent genera, but the 
differences are obviously much more dramatic under the binomial system of 
nomenclature if three genera are recognized rather than one genus. 

Unfortunately the International Code of Botanical Nomenclature (ICBN 
1 988) still provides remarkably little guidance in the matter of choosing a 
lectotype. In the absence of clear directions from ICBN, botanists will of 
necessity flounder along with their divergent conclusions as to what the lec- 
totype of such Linnean genera as Myrica should be. A special Committee on 
Lectotypification was established by the Nomenclature Section of the Sydney 
Congress to resolve problems such as those pointed out by Stirton et al. 
(1981). This Committee recommended to the Berlin Congress that Art. 8.1 
be amended to read "The author who, on or after 1 January 1 93 5 , first desig- 
nates a lectotype or a neotype must be followed. ..." It was hoped that this 
stipulation would eliminate the uncertainty connected with use of the term 
"type" by such early authors as Rafinesque whose concept of type surely was 

different than that of a present-day investigator and also the uncertainty of 
the lectotypes designated under both the American Code and the Type Basis 
Code. Under this suggested requirement, the genus Myrica would have been 
typified by M. gale as that was the choice of Hitchcock and Green and pub- 
lished as an unofficial supplement to the ICBN ( 1 93 5 , p. 1 1 6). However, the 
International Botanical Congress meeting in Berlin in 1988 found itself 
unable to resolve the problems of lectotypification and, since one committee 
failed to solve the problem to everyone's satisfaction, three committees were 
appointed to study and to report their recommendations to the 1993 Con- 
gress (Tokyo). As might be expected the complicated problems of lectotypi- 
fication have been largely passed on to the next Congress. 

Both Myrica gale and M. cerifera have been designated as the lectotype for 
the generic name Myrica. The species best known to Linnaeus as the only mem- 
ber of the genus occurring in Sweden, where it is abundant, is Myrica gale. 
Britton, operating under the American Code, designated this species as the 
type (= lectotype) of the genus. Hitchcock & Green (1929) made the same 
choice; their conclusions as to the "standard species" (= lectotype) were 
listed in the International Rules (1935) in a semi-official way. However, the 
desirability oi Myrica gale 2iS a lectotype was challenged by Hylander (1945) 
since that species with a very few close allies had been segregated as a small, 
independent genus. If the much larger clade, represented by Myrica cerifera, 
were to be segregated from Myrica this larger portion of the genus, a group 
of ca. 50 species, would belong to this segregate almost all requiring new 
combinations. Rehder (1949) also accepted Myrica cerifera as the lectotype 
of the genus Myrica. In spite of this consideration, it seems certain that 
Myrica gale, the historically best or at least the scientifically longest known 
species o( Myrica, will be confirmed as the lectotype of the genus. Prelimi- 
nary reports of the subcommittee dealing with lectotypification of Linnaean 
generic names strongly suggest that Myrica gale will be recognized as the 
lectotype o{ Myrica L. 


herital n 


rical nutlet cc 

)vered by 

fruit lack 


ent; dis 

tal axillar 

y buds flora 

1; mature 


pillae, somewhat 




• flattened 

and keele. 


Ileal to cylmclnc. 



•d nor papilk 


Leaves stipulate 


flowers 2, unlob 

ed, adm 

ite to the 

fruit; fruit a 


keeled nu 

rendered buoyan 



, enlarged, un 





2. Leaves stipulate; leaf margins deeply pinnatifid wkh broadly rounded 
lobes; bracteoles of pistillate flowers 2, deeply lobed into linear segments, 

enveloped at maturity in a bur-like involucre formed from the lobed 
bracteoles and the accompanymg bracteal scale 3. Comptot 


Morella Lour., Fl. Cochinch. 548. 1790. Type: Morella rubra Lour. 

1. Aments branched; pistillate bracts subtending several ovaries of which only 
one develops; fruit 6-8 mm in diameter, covered at maturity by imbricate, 
fleshy, succulent papillae subg. 1. Morella 

I. Aments simple; pistillate bracts usually solitary, simple, and l-flowered; 

neither fleshy nor succulent papillae subg. 2. Cerothamnus 

Morella Lour., FL Cochinch. 548. 1790. Type: Morella rubra Lour. Mynca sect. 
Morella (Lour.) Benth. & Hook.f., Gen. PI. 3:401. 1880. Mynca subg. Morella (Lour.) 
Engler, Nat. Pflanzenfam. IL 1:27. 1893. 
Subgenus Morella apparently contains fewer than 10 species of eastern 
Asia, the Philippines, and Malaysia. The differences between the species of 
subg. Morella and subg. Cerothamnus from both the Americas and central and 
southern Africa are so striking as to have elicited expressions of consterna- 
tion from Greene (1910) that Small ( 1 903) would transfer our southeastern 
species to Morella; this dismay was shared by Nieuwiand (1910). Greene 
could not believe that plants whose fruits were so palatable and wholesome 
as to be eaten both uncooked and cooked and so succulent and juicy as to be 

fruited plants of the southeastern United States or the western coastal region 
from Vancouver Island throughout much of California. Perhaps Greene was 
correct in his assessment, but for the present, based largely upon the findings 
but not the conclusions of MacDonald (1978) and Abbe (1972), the Asiatic 
plants are here treated as merely subgenerically differentiated from their 
American and African congeners. 

Morella Subgenus Cerothamnus (Tidestr.) Wilbur, comb. & stat. nov. 

Cerothamnus Tidestt., Elys. Marian., Ferns. 41. 1910. Lectotype: Cerothamnus arborescens 
(Castigl.) Tidestr. (= Myrka cerifera L.) [herein designated}. 

Usually aromatic, dioecious or polygamo-monoecious shrubs to small 

evergreen, entire or serrate, estipulate. Inflorescences borne proximally be- 
low or axillary to the lower leaves, the staminate erect or nearly so, at anthesis 

thick-cylindric to broadly ellipsoid, bracts broadly to narrowly ovate and at 
anthesis shorter than the 1-22 stamens, the stamens yellow or becoming 
yellow, filaments simple or branching and arising from the staminal column 
at different levels, the secondary, tertiary and even quarternary bracts often 
present; pistillate inflorescences simple or basally branched, at anthesis 
ovoid to cylindrical, the rachis sometimes glandular and the bracts usually 
persistent, the pistillate flower subtended by secondary, tertiary, or even 
quarternary bracts forming a calyculus, the ovary either glabrous or pilose 
and almost completely covered by persistent, more or less globular, wax- 
secreting papillae. Fruit a nutlet, ± spherical and mostly covered by a layer 
of wax, the ovary wall glabrous to densely pilose, the papillae glabrous to 
puberulent, the associated secondary, tertiary, and even quarternary bracts, 
if all present, persisting until after fruit maturation and never enlarging. 

Cerothamnus, the largest subgenus in Myricaceae, contains the waxy- 
fruited species of Morella native to the Americas and Africa and their 
neighboring islands — i.e. the West Indies and the Atlantic islands lying off 
the northwestern coast of Africa. 

:; fruit wall 

Morella series Cerothamnus (Tidestr.) Wilbur, comb. & stat. nov. 

Cerophora subg. Cerocarpa Raf., Alsogr. Amer. 11. 1838. Lectotype: Cerophora lanceolata 

Raf., herein designated {= Myrica certfera L.] 
Myrka sect. Cerophora (Raf.) A. Chev., Mem. Soc. Sci. Nat. Cherbourg 32:223 (= Monogr. 

Myric. p.l39) 1901. 
Cerothamnus Tidestr., Elys. Marian., Ferns. 41. 1910. Lectotype: Cerothamnus arborescens 

(Castigl.) Tidestrom (= Myrka certfera L.) 
Aments simple; pistillate usually solitary, simple, and 1 -flowered. Fruit 
1-5 mm in diameter. Papillae wax-producing, neither juicily succulent nor 

In spite of the few species in Morella series Cerothamnus present in eastern 
North America, there currently exists a surprising amount of uncertainty as 
to just how few or how many species can be recognized in that well-collected 
area. The number of species are in any event few: perhaps no more than two 
and certainly no more than four. 

2. A second taxon has been proposed whose distinctness from Morella cerifera 
is denied by several of our more experienced students of the southeast- 
ern flora. The questioned taxon is usually a low bushy, rhizomatose 
plant with strikingly smaller leaves. It was treated by Michaux (1803) 
and Radford et al. (1 968) as a variety {Myrka cerifera v^t.putmla Michx.) 
but as a species by Rafinesque (1838) and Small (1903 and 1933) 
{Myrica pusilla Raf., Morella pumila (Michx.) Small, and Cerothamnus 
pumtlus (Michx.) Small). 
Many field observers and collectors of bay berries, are at least partly 
convinced that Myrica pusilla merits taxonomic recognition for these dwarf 
plants seem strikingly unlike the much more abundant and luxuriant M. 
cerifera that often grow in close proximity. In the herbarium on the other 
hand the claim to specific or even varietal status of M. pusilla seems much 
less certain perhaps because most specimens are not accompanied by notes 
as to either the height or habit of the plant; those two features together with 
the much smaller size of the leaves are what makes these plants in the field 
so strikingly unlike the ubiquitous M. cerifera. In spite of the apparent dis- 
tinctiveness noted in the field, it seems impossible to maintain as distinct 
cerifera and pusilla even in a group as plastic or as bereft of taxonomically 
useful characters as are the bayberries. Sufficient dissatisfaction with this 
conclusion exists, however, that I intend and would urge all other botanists 
with the opportunity to study and collect the plants in the field to do so and 
to record carefully notes on the height, habit and habitat of the plants — 
something that surprising has been largely neglected by most collectors 

Thieret (1966), who has had extensive field experience with both species 
in Louisiana, noted that "from a clump that is otherwise typically M. pusilla, 
there will arise one main stem to ten or twelve feet tall and six inches in 
diameter — a M. cerifera stem. Examination reveals that all the stems of the 
clump comprise a clone. . . . Plants intermediate in habitat between the two 
extremes are usually found in the vicinity." Thieret concluded that "the 
habit differences ... are not reliable criteria but are simply responses to 
habitat differences" and "that the habit extremes pass insensibly into each 
other." I too have often observed in North Carolina both pusilla and cerifera 
growing in close proximity but have not yet noted the blurring of growth 
form and leaf size that Thieret has noted nor am I convinced that there is 
always a correlation of habitat and growth form. I have regrettably never 
tried to excavate the underground connection perhaps implied by Thieret 
and which, if proven, would provide convincing support for the treatment 
tentatively accepted here of non-recognition (unless root grafting occurred). 
Godfrey (1988), whose field experience with our southeastern plants is 

surely unrivalled, does not accept the dwarf plants as a taxon deserving a 
name either. Additional study is certainly needed, especially investigations 
centered in the field. 

^ill.) Small, Fl. SE U.S. 337 & 1329. 1903 [as 

Loisel. as usually < 

Although my study is not completed, I am doubtful that the above two 
species can be maintained as distinct. The alleged principal difference 
between the two is apparently to be found in the fruits: fruit of more 
northern species {Myrka pensylvanka) possess puberulence on both the 
papillae and ovary wall; fruit of the southern species (usually referred to as 
Myrka heterophylla Raf.) have glabrous papillae and glabrous walls. Due to 
the dense covering of wax, detection of the puberulence is often difficult and 
the amount varies from a dense covering to very few trichomes. The other 
alleged differences of fruit size, twig pubescence, leaf persistence, etc. seem 
even less consistently diagnostic. If only one species is to be recognized, the 
correct binomial is Morella carolinknsis (Mill.) Small, a binomial that has 
been applied to the combined species in the past and to both of the species 
at different times when they were treated as distinct species. Miller's name 
should be applied to the species with the more southern distribution if two 
species are represented as most authors have accepted for the past 75 years. 

5. Morella inodora (W. Bartram) Small, Fl. SE U.S. 337 & 1329. 1903. 

Myrica inodora W. Bartram, Travels Carolina, 405. 1791. Cerothamnus inodorm (W. 

Bartram) Small, Florida Trees 12 & 102. 1913. 
In striking contrast to most other Myricaceae, but as indicated by its 
specific epithet, the crushed foliage q{ Morella inodora is not aromatic. The 
species ranges from southern Georgia westward into southeastern Louisiana. 

6. Morella californica (Cham. & Schltdl.) Wilbur, comb. nov. Kyrua 

califormca Cham. & Schltdl., Lmnaea 6:535. 1835 . Galecalifornka (Cham. & Schltdl.) 
Greene, Man. Bot. San Francisco. 298. 1894. 

The natural range of this speci( 
Washington south into Los Angef 

MORELLA series FAYA (P. Webb & Berthel.) Wilbur, comb. & star. nov. 

Fayana Raf., Alsogr. Amer. 12. 1838. Type: Fayana azonca Raf. (= Myrica faya Alton). 

Faya P. Webb & Berthel., Hist. Nat. lies Canaries 3:272, t. 216. 1847. Type: Faya 

fragifera P. Webb & Berthel. (= Myrica faya Alton). Myrica sect. Faya (P. Webb & 

Aments simple or branched; pistillate bracts subtending several ovaries 
of which only some usually develop. Fruits usually A-6 mm in diameter, 
often forming a syncarpium. Papillae often producing wax but never fleshy. 

Series Faya is a small taxon of three geographically widely separated 
species. Besides the two North American representatives treated below, the 
section is composed of the type species of the section, Morella faya^ of the 
Canaries, Madeira, and the Azores and possibly also of Portugal where it 
occurs but perhaps only as a naturalized introduction Burges (1964). 
MacDonald (1977, p. 2638), who has presented a series of papers on the 
morphology of the inflorescence of many myricaceous taxa, is of the opinion 
that "section Faya could easily be incorporated in section Cerophora" of the 
genus Myrka. This genus in his opinion would then consist of the fleshy- 
fruited section Morella of eastern Asia and Indonesia and section Cerophora^ 
the largest taxon in the family, with both American and southern African 
representatives. The suspicion lingers that series Faya is not a proven 
monophyletic group not only because MacDonald questioned its morpho- 
logical distinctiveness but also because its distribution pattern is not one 
readily explained or matched by other examples. 

In contrast to the uncertainty existing in both the identities and names 
of the taxa comprising Morella series Cerothamnus, the taxa forming series 
Faya are morphologially most distinct and geographically widely separated . 

Myrica L., Sp. PI. 1024. 1753; Gen. PL ed. 5.449. 1754. Lectotype: Mynca 


imort.,Fl.Belg. 12.1827 

. [Neither Gale 



•br.Arbust. 1:25 

1 7 5 5 ) nor Gale Adanson . , (Fam 


1 Duhamel's and Adanson's reintrodu 


of Ga/e were illegitimate names as su 


tions for My. 

nca L. and hen 


;nclaturally superfluous {J 

Vrt. 63 ICBN)]. 


"b" Gale tTourn.] Endl., 

Gen. PI. 272. 


{Employed h 

1 the sense of t 


caceae less Comptoma.) 


ra subg. Galesm Raf., Alsogr. Amer. 11. 


, Lectotype: ( 

Zerophora {Galest 


:f^fonaRd.(= Myrica gale 


Angeta ' 

rns37. 1910. Type: A 

ngeia palmtris 



capaius'ms-Um., = Myrm 


). nov. BASIONYM: Mynca fay 

Aromatic, usually dioecious shrubs lacking terminal buds. Leaves alternate, 
pinnately veined, serrate to entire, deciduous, estipulate. Inflorescences 
borne distally, the staminate suberect to recurved, in bud ovoid to elliptica 
and at anthesis cylindrical, the bracts broadly ovate to triangular, at anthesi 
longer than the 3-6 stamens, secondary bracts absent; pistillate inflores 
cences simple, broadly cylindric at anthesis, the bracts persistent, the pis- 
tillate flowers each subtended by 2 secondary bracts partially adnate to the 
ovary wall, the ovary glabrous and lacking papillae. Nutlet flattened, keeled, 
not covered with wax but with few to many glandular trichomes, the brac- 
teoles persistent, becoming greatly enlarged and inflated, strongly adherent 
to the flattened fruit, inflated and forming a buoyant float. 

Myrka is represented in the Americas by two species that are morphologi- 
cally readily distinguished from one another and whose geographical ranges 
do not overlap. 

1. Myrica gale L. 

This species has a broken circumboreal distribution pattern extending 
) northern New Jersey, Pennsylvania, 
Lakes from as far west as eastern 
Minnesota and in the west to the mountains of Oregon. Morphological 
variation apparently correlated with distribution occurs within the exten- 
sive range of this species but its taxonomic merit requires additional study. 
Hulten (1944 and 1968) presented a brief overview of the problem. 

2. Myrica hartwegii S. Watson. 

This species is found along stream banks in yellow pine forests in the 
Californian Sierra between 300-1500 m and consequently not sharing any 
part of its range with any other member of the Myricaceae. 

3. COMPTONIA UHer. ex Alton 
Comptonia L'Her. ex Alton, Hort. Kew. 3:334. 1789- Type: Comptoma 

asplemfolia (L.) UHer. ex Alton (= Comptonia peregrina (L.) J.M. Coulter). 

Myrtca "c" Comptoma (L'Her. ex Alton) Endl., Gen. PI. 272. 1837. 

Mynca sect. Comptoma (L'Her. ex Alton) Endl. ex CDC, Prodr. 16(2):151. 1864. 

Myrica subg, Comptoma (L'Her. ex Alton) Engler, Nat. Pflanzenfam. III. 1:28. 1893. 

Aromatic, usually dioecious, colonial shrubs 1.5m tall or less and lacking 
terminal buds. Leaves alternate, pinnately veined and lobed, deciduous, 
conspicuously semi-cordately stipulate. Inflorescences borne distally, the 
staminate suberect to strongly recurved, cylindrical, the primary bracts 
persistent, broadly ovate to quadrangular, at anthesis longer than the 3-7 
stamens, adaxally bearing numerous glandular trichomes, secondary bracts 

absent; pistillate inflorescences simple, broadly ovoid at anthesis, the rachis 
pubescent and glandular, the bracts persistent but in fruit obscured by the 
greatly enlarged lobes of the secondary bracts, the pistillate flower sub- 
tended by the 2 greatly enlarged and deeply lobed secondary bracts, the 
ovary glabrous and lacking papillae. Nutlet conical to cylindrical, non- 
ceriferous, 2.2-5.5 mm long, glabrous, surrounded by the enlarged lobes of 
the secondary bracts, which form a bur-like structure in fruit. 

The genus Comptoma is both monotypic and endemic to eastern North 

1. Comptoniaperegrina(L.)J.M. Coulter, Mem. Torrey Bot. Club 5:127. 
1894. Ltqmdambar peregrtna L., Sp. PI. 999- 1753. Myrica peregrtna (L.) Kunrze, 
Revis. Gen. PI. 2:638. 1891. 

Mynca asplenifolia L., Sp. PI. 1024. 1753. Ltqmdambar asplemfolia (L.) C.F. Ludwig, 
Neuere Wilde Baumz. 27. 1783. Comptonia asplenifolia (L.)i:H6r. ex Aiton,Hort. Kew. 
3:334. 1789. Comptoma peregrma var. asplenifolia (L.) Fernald, Rhodora 40:4l0. 1938. 

The variation in vegetative pubescence suggested by Fernald as a diagnos- 
tic feature distinguishing \a.r. peregrtna from var. asplenifolia (L.) Fernald does 
not delimit populations as sharply as Fernald's account implied. There seems 
to be more continuous variation in pubescence than Fernald reported and the 
geographic range of the two pubescence types is less discrete than suggested. 
Well-collected areas of the mid- Atlantic States have many examples of both 
pubescence types within their borders . Examples of different pubescence types 
on different branches of the same shrub are not unusual. It surely can be con- 
cluded that varietal status within the monotypic genus Comptonia based on 
pubescence has not been convincingly proven. My rapid survey of hundreds 
of specimens does not suggest that a detailed, careful analysis of pubescence 
would likely demonstrate the presence of geographically based varieties. 

As shown by the above synonymy, Linnaeus treated this species twice in 
Species Plantarum. The species was these first included (p. 999) as Liquidam- 
barperegrina and again (p. 1024) as Myrica asplenifolia. For nearly a century 
and a half the name most frequently adopted was Myrica asplenifolia or its 
derivative Comptonia asplenifolia. For the past century (except for Gleason 
(1952), Gleason&Cronquist (1963), and Wagner etal. (1990), the accepted 
name has been almost universally that based upon Liquidambar peregrina. 
The latter choice is mandated by the International Code not because of the 
American Code's favoring of the name appearing first in a volume ("page 
priority") but because Linnaeus, upon discovering the conspecificity of the 
two binomials, was the first to unite them (Syst. Nat. ed. 10.2: 1273.1759.) 
by placing Myrica asplenifolia in the synonym oi Liquidambar peregrina. 

jRANDEGEt, K. 1901. Some sources of error in genera and species. Zee 5:91-98. 
JuRGES, N.A. 1964. Myricaceae in T.G. Turin et al. Flora Europaea. 1:56. 
Ihevalier, a. 1901. Monographie des Myricacees; anatomic et histologie, organo, 

classificarion, et description des especes, distribution geographique. Mem. Soc. J 

Cherbourg 32:85-340. 

. Nat. Pflanzenfam. 
manual of botany. I 

1952. The new Bntton and Brc 

Gleason, H.A. and A. Cronquist. 

Gleason, H.A. and A. Cronquist. 1991. Manual of vascular plants of northeastern United 
States and adjacent Canada. 2nd edition. New York Botanical Garden, Bronx. [Myricaceae, 

Godfrey, R.K. 1988. Trees, shrubs and woody vmes of northern Florida and adjacent 
Georgia and Alabama. Univ. of Georgia Press. Athens. {Myricaceae, 483-489.] 

Greene, E.L. 1910. Nomenclature of the bayberries. Leafl. Bot. Observ. Crit. 2:37-40. 

Greene, E.L, 1910a. A new name for the bayberries. Leafl. Bor. Observ. Crit. 2:101-104. 

Hftchcock, A.S. & M.L. Green. 1929. Standard-species of Linnean genera of Phanerogamae 
(1753-54.) International Bot. Congress, Cambridge, 1930 inNomencl. Prop. Brit. Bot. 
1 10-199. His Majesty's Stationery London Office. 

Hulten, E. 1944. Flora of Alaska and the Yukon. ActaUniv. Lund. Avd. 2.40: 572 [Myrwa]. 

1968. Flora of Alaska and neighboring territories. Stanford University Press, 

Stanford, CA. [Myrica p. 364}. 
luTCHiNSON, J. 1964. The genera of flowering plants. Dicotyledones. Oxford Univ. Press. 

London [Myricaceae, 2:120-121. 1967.} 
Iylander, N. 1945. Nomenklatorische und systematische Studien iiber Nordische 

Gefasspflanzen. Uppsala Univ. Arsskrift. 7:1-337. [Myma, 40.} 
MGER, R.F. 1958. Comments on the definition of genera. Evolution 12:370-384. 


Fischer, Jena. 

Regnum Veg. 97:xiv, 1-457. Bohn, Scheltema & Holkema. Utrecht. 
1988. "Berlin Code." Regnum Veg. 1 1 8:xiv & 328. Koeltz Scientific Books. 

Konigstein, Germany. 
Iacdonald, A.D. 1977. Myricaceae: floral hypothesis for Gale and Comptonia. Canad. J. 

Bot. 55:2636-2651. 




:,A. 1 








lerches sur la classification des chenes. Bianco Luno-F.S. Muhle, 

Radford, A.E., H.E. Ahles and C.R. Bell. 1968. Manual of the vascular flora of the 

Carolmas. Univ. North Carolina Press, Chapel Hill. {Myricaceae, 360-361.] 
Raeinesque, C.S. 1838. Alsographia americana. Philadelphia. [Myricaceae, 8-12.} 
Rehder, a. 1949. Bibliography of cultivated trees and shrubs. Arnold Arbor. Harvard 

Univ., Jamaica Plain, MA {Myricaceae, 87-88.] 
Rendle, A.B. 1903. Notes on Myricaceae. J. Bot. 41:82-87. 
ScHWARZ, O. 1936. Entwurf zu einen Natiirlichen System derCupuiiferen und der Gattung 

QuercusL. Notizbi. Bot. Gart. Berlin-Dahlem 13:1-22. 
Small, J.K. 1903. Flora of the southeastern United States. New York. [Myricaceae, 


1933. Manual of the southeastern flora. New York. {Myricaceae, 408-410.] 

Stirton, C.H., D.V. Field, R.K. BRUMMiTxandJ. McNeill. 1981. Proposals on stability of 

names. Taxon 30:250-256. 
Thieret, J.W. 1966. Habit variation in Mynca pensylvamca and M. cerifera. Castanea 


Wagner, W.L., D.R. Herbst and S.H. Sohmer. 1990. Manual of the flowering plants of 
Hawaii. 2 vol. Univ. Hawaii Press. {Myricaceae 1: 929-931.] 


Ramamoorthy, T.R, R. Bye, A. Lott and J. Fa (Eds). 1993. Biological 
Diversity of Mexico: Origins and Distribution. (ISBN 0- 1 9-5 06674- 
X, hbk) Oxford University Press, New York. $79.95. 812 pp. 

expertise, they are at the same time likely to rely on the volume as an invaluable source of 
data and bibliographic references on the subject of Mexican biodiversity. 

And the subject is an extremely important one. Although Mexico is the fourteenth largest 
country in the world, it is usually ranked within the top three or four countries in the world 
in terms of the richness of its biological diversity. This volume does a very good job at 
explaining, in general and taxon by taxon, how biogeography, geology, and evolutionary 
history have combined to bring about Mexico's fantastic species richness. If the book does 
nothing more than draw well-deserved conservation attention to Mexico's arid and montane 
habitats, it will have performed a great service. 

The book's 26 chapters are organized into six sections. The first section's three chapters 
focus on historical background, with chapters on geology and the origins and diversity of 
Mexico's flowering plant flora. The second section reviews six selected faunistic groups 
including: bees, butterflies, fishes, herps, birds, and mammals. A chapter on ecological 

reviews 1 1 floristic groups including pteridophytes, bryophytes, grasses, and legumes. The 
two chapters covering the genus Ptnus and Quercus are particularly outstanding. The fourth 
section contains two chapters analyzing phytogeographical patterns in contrasting ecosys- 
tems: tropical rainforests and alpine habitats. The fifth section contains two chapters dealing 
with ethnobotanical themes, which although unlikely to contain much information that is 
new to those familiar with the field, provide a compilation of bibliographic references that 
is very useful. The sixth and final section contains an overview of Mexican biodiversity in 

Some of the chapters are quite noticeably dated; the symposium which was the inspiration 
for the present volume was held in 1988. Other chapters have been recently updated, and, 
in general, the lack of bibliographic references from the early '90s is a minor drawback, and 
one that is probably unavoidable in a volume of this scope. There are translation problems 

chapter in which the translation seems particularly weak is that by Hernandez-X. on plant 

Many of the chapters contain conservation recommendations at the end which although 
skeletal, do in some cases (e.g., the chapter on Pinus) make specific important recommen- 
dations. However, it is not this volume's place or role to produce comprehensive conserva- 
tion recommendations, but hopefully to provoke the development of these by others. 



Departamento de Botdmca 

JJniversidad Autonoma Agraria "Antonio Narro" 

Buenavista, Saltillo 

Coahuila 23513, MEXICO 

es seco. Se eleva desde los 1200 m hasta los 2350 m. Las rocas que la forman : 
fundamentalmente calizas y los suelos litosoles. No presenta escurrimentos permanentes 

Poaceae y Fabaceae, los generos Muhknbergia, Daiea, Notholaena y Polygala presentar 

substrate and lithosoles the principal soils. There are no permanent water : 
The vegetation is divided in four types: Chihuahuan Desert Scrub, Submoi 
pine Woodland and Grassland. The flora comprises approximately 87 i 

La sierra de la Paila es una formacion montafiosa aislada que forma p 
del complejo de la Sierra Madre Oriental. Se localiza en el limite este 
Desierto Chiuahuense y es un mosaico de asociaciones vegetales que cam 

ir ser un area poco estudiada botanic 
■ibuir al conocimiento de la flora del n. 

La Sierra 

de la Paih 





la forma r 



. Tiene ap 


5n montanosa domica aislada, ubicada 
g. 1 ). El area ocupadapor la sierra tiene 
■elos 101°25'y 101°49'Wylos25°45' 
; unos 60 km de extension en su eje 
mayor (norte-sur) y unos 35 km en su eje menor, con una superficie de unos 
1700 km2. Se eleva desde los 1200 m alcanzando las partes mas altas los 
2350 m de altura. Esta surcada por una serie de cafiones que van del centre 
a la periferia. En la parte central se localizan valles altos y pequeiios. 

Varias vias de comunicacion bordean la sierra totalmente. Por el sur pasa 
la carretera 40 y la linea de ferrocarril Saltillo-Torreon. Por el noreste pasa 
la carretera 57 y el resto de la circunferencia los forman caminos de terracerfa. 
El poblado de Hipolito, ubicado en el extremo sur, es el mas importante y 
proximo a la sierra. El ejido Las Coloradas se localiza en los valles cercanos 
al extremo norte. Los ejidos de El Cedral y Parreiios se encuentran dentro de 
la sierra (Fig. 2). 

De acuerdo a lo presentado por el INEGI (1983), los climas seco y 
semiseco (BS°,BSi) dominan en el area. Estos climas se caracterizan por 
presentar Uuvias predominantes en el verano y temperaturas altas. La 
precipitacion promedio es de 400 mm anuales, y la temperatura media anual 
es de 20°C. 

Las rocas dominantes en el area son las calizas, en su mayoriadel Cretacico. 
Los suelos predominantes son litosoles, arcillosos y ricos en materia organica, 
de color pardo oscuro o negro. Los valles intermontanos de la sierra presentan 
xerosoles, y las partes bajas y valles que rodean la sierra presentan suelos 
aluviales tipo castanozem y ferozem (INEGI 1983). 

En la sierra no se presentan correientes permanentes ni depositos de agua, 
solo arroyos en los caiiones que drenan el agua durante la epoca de lluvias. 

Aunque la colecta de muestras vegetales en la Sierra de la Paila se 
realizaron desde principios de siglo (Sousa 1969), la informacion sobre la 
vegetacion del area es escasa. En trabajos tradicionales como los de Muller 
(1947), pasa desapercibida. Solo recientemente se ha aportado informacion 
por Cano y Marroquin (1967) y Wehbe (1985). 

La vegetacion en el area de la Sierra de la Paila puede ser clasificada en 
cuatro tipos prnicipales, que de acuerdo con la revision sobre vegetacion de 
Coahuila hecha por Villarreal y Vlades (1992-1993) correspeonden a: 
Matorral Desertico Chihuahuense, Matorral Submontano, Bosque de Encmo- 
pino y Zacatal. 

Matorral Desertico Chihuahuense. Es el tipo de vegetacion mas comu 
en coda el area, circundando coda la sierra. Los valles que bordean la sierra 
lomerios con suelos algo profundos y poco pedregosos son ocupados pc 
macorrales microfilos de arbuscos de 30-150 cm de ako. La especie m: 
frecuence es Larrea tridentata, asociada con Flourensia cernua, Fouquien 
splendens, Parthenium incanum. Yucca filif era, Y . treculeana, Opuntia imbrkata 
Prosopis glandulosa. En algunos sicios de la porcion oesce Serkodes greggii ( 
frecuence y Grusonia hradtiana abunda en las porciones norce. 

Las laderas de las pociones mas excernas de la sierra, asi como los lados mi 
expuestos de muchos cafiones con suelos pedregosos y someros se desarrolla 

matorrales bajos con comunidades de Agave lechuguilla, asociada 
frecLientemente con Parthenium argentatum, Euphorbia antisyphilttica y con 
presenciade A^^z^ej/f/^to, Dasylirionpalmeri, Hechtia texensh, Opuntia microcLtsys, 
0. phaeacantha. Yucca earner osana y Gochnatia hypoleuca. 

Matorral Submontane. Este tipo de vegetacion ciene distribucion 
irregular, se localiza en los canones y usualmente se mezcla con porciones de 
bosque en el centro de la sierra. Esta formado por arbustos y pequenos arboles 
de 2-5 m que se desarroUan en comunidades densas en los lechos de los 
arroyos y areas protegidas con suelos profundos y algo de humedad. Se 
presentan frecuentemente Quercus mvaginata, Q. intricata, Fraxinus greggii, 
Vauquelinia corymbosa, Cercocarpus mojadensh. Pis facta texana. Acacia berland- 

VILLARREAL Q., Flora Vascular de la Sierra de la Paila 113 

leri, Rhus virens, Flourensia retinophylla, Berberis trifoliolata, Ptelea trifoliata y 
Leucaena greggiL Brahea berlandieri crece en las laderas de los cafiones. En los 
lechos de los arroyos es frecuente Dodonaea viscosa. 

Bosque de Encino-Pino. Se localiza en la porcion central de la sierra, 
ocupa valles y canones amplios con comunidades arboreas poco densas do- 
minadas por Quercus graves ii, Q. laceyijuniperus flaccida, con elementos espar- 
cidos de Pinus arizonka o P. cembroides. Otras especies frecuentes incluyen a 
Juniperus erythrocarpa. Arbutus xalapensis, Garrya ovata, Ungnadia speciosa, 
Fraxinus cuspidata y elementos de matorral submontane. Proximo al bosque 
es frecuente encontrarvegetacionarbustivadetipo chaparral, que usualmente 
se mezcla con las especies arboreas del bosque. En estas comunidades esta 
presente Quercus intrkata, Q. invaginata, Q. grisea y Rhus virens. En los arroyos 
es ft:ecuente J uglans microcarpa. Las especies arboreas usualmente presentan 
especies de Til lands ia y P horadendron . 

Zacatal. En areas reducidas de los valles centrales elevados de la sierra con 
suelos profundos se propicia el desarroUo de comunidades de gramineas. En 
dichos zacatales sobresalen Sporobolus airoides, Nassela tenuissima, Stipa eminens 
y en otras areas Bouteloua gracilis y B. curtipendula, frecuentemente mezcaldas 

La flora vascular documentada de la Sierra de la Paila comprende 703 
especies (712 taxa incluyendo a los taxa infrespecificos) distribuidas en unos 
387 generos y unas 87 familias. Es ia localidad tipo de unas 21 especies, de 
las cuales 9 son endemicas (ver tablas 1 y 3). Las familias y generos con mayor 
diversidad se muestran en la tabla 2. 

Las primeras colectas parecen haber sido hechas por Endlich en 1 905 y C. 
A. Purpus enl910yl911, cuyo material sirvio para la descripcion de nuevas 
especies. En los ultimos afios una docena de colectores ha trabajado en la 
sierra, como se presenta en la lista de abreviaciones. Los herbarios revisados 
para la elaboracion de la lista floristica fueron el de la Universidad Atomoma 
Agraria Antonio Narro (ANSM) y el de la Universidad de Texas (TEX-LL). 

El arreglo que se sigue para la lista de plantas vasculares en los grupos 
principales es el de Cronquist ( 1 98 1 ) para angiospermas y el de Crabbe et al. 
(1975) para helechos y grupos afines a helechos. Las categorias de familia, 
genero y especie se presentan en secuencia alfabetica. Cada nombre cientifico 
va seguido por el tipo de vegetacion donde frecuentemente se encuentra, 
luego los datos sobre el habitat y frecuencia. Al final la abreaviacion de los 


Flourensia retinophylla Blake Poaceae 
Stevta saliafoha Cav. var Integra (Blake) Rob. Bou, 

*EMnocereus delaetii (Giierke) Giierke Rha 

*Gymnocactus aguirreanus Glass & Foster Rubiaceai 


Tipos de vegetacion: MD, Matorral desertico chihuahuense. MS, Matorral 
submontano. B, Bosque de encino-pino. P, Zacatal. Principales colectores y 
herbarios donde estan depositados sus colecciones: C, M. A. Carranza P., 
(ANSM). Ca, G. Cano yj. S. Marroquin (UNL). CQ, D. Castillo Q., (ANSM). 
Endlich, R. Endlich (MO). H, J. Henrickson (UC, TEX-LL). Hin, G. B. 
Hinton (TEX-LL). JM, J. S. Marroquin (ANSM). JW, J. Wehbe (ANSM). 
MCJ, M. C. Johnston, E Chiang y T. Wendt (TEX-LL). Purp, E A. Purpus 
(UC,F,US). R, A. Rodriguez G. (ANSM). VR, J. Valdes Reyna (ANSM) y V, 
J. A. Villarreal Q. (ANSM). 





ng. MS. Paredes rocosas. Comun. V3L 




..Br. MS, MD. Paredes 

rocosas. Escasa. V; 

J094, 3559, 5312. 


wrightii ] 

riieron. MD, MS. Lade, 

ras rocosas. Comun 

.V3095, 3533, 441 






;^/^^^;.2m«(Buckl.)Kunt2e. B 

. Canones y arroyos 

;. Escasa. V3524, 4 




lotdes (Desv.) Maxon. B, 

MS. Canones y am 

oyos. Comun. V31( 


4479, 4730, 528 

8, VR2233. 

Cheilanthes pinkavae 

Reeves. MD. Laderas j 

-ocosas. Escasa. V3' 



t aschenbor 

■mana Kl. MS, MD, B. 

Laderas rocosas. Cc 






; bryopoda 

Maxon. MD, MS. Lac 

leras y arroyos. Co 

mun. H16543, MCJI0099, 

10109, 10511, R1486. 
Notholaena cochtsensts Goodd. MD. Laderas rocosas. Comiin. H16512, CQ783, V4727. 
Notholaena greggii (Mett.) Maxon. MD. Laderas rocosas. Rara. MCJ 105 20, V5562. 
Notholaena integerrima (Hook.) Hevly. MD. Laderas rocosas. Escasa. CQ782, 787, R1481. 
Notholaena neglecta Maxon. MD. Laderas rocosas. Escasa. MCJl 171 le, R794, V5201 , 5259, 

Notholaena parvifolia Tryon. MD, MS, B. Canones y arroyos. Comun. MCJ10109b, CQ660, 

V3098, 3542,4472. 
Notholaena sinuata (Lag.) Kaulf. MD, MS. Laderas rocosas. Comun. V3097, 3591. 
Notholaena standleyi Maxon. MS. Laderas rocosas. Escasa. V4480. 
Pellaea atropurpurea (L.) Link. B, MS. Canones y arroyos. Escasa. V3523, 5277, 6660, 



PINOPHYTA (Gimnosper 

laderas. Comun. V3561. 


Ephedra anUsyphtlittca Mey. MD. Laderas. Escasa. JM2327, MCJ10512, 10519, V2892, 

Ephedra aspera Wats. MD. Laderas. Escasa. JM320, C462, V3873, 5572. 

MAGNOLIOPHYTA (Angiospermas) 
MAGNOLIOPSIDA (Dicotiledoneas) 



mhus hma 

yos \ can 

.ones Co 


'220, MCJl 





//w/„ Hiaiidi; \Il^ Ari(i\<)s ( omun l^irp r5 

1, V5436 

r.ghtiu .upi 



(-Uv(.)Kunt/L MS, U, MD, P ( omL'm CQ- 

10, V390 



parryi Gra' 

, MD \ lade. as Mu> comi.n CQ6^5 

, MCJl 16 

90a, HK 

Siphomglos.a ptlosei 

V.^(nes)'Io.. MD, iVLS Valks) Es.asa 






'//MIook MD Vallc-s Es.asa CQ8()6. 






Tho.nb. MS, MD. Arroyos. Comun. H 16187, 

V3902, ^ 

Tukstromia lanugmoui (Nutt.) Standi. MD, 1 

Tidestrotnta suffrutkosa (Torr.) Standi, var. coak 

Purp4927, H16195, V2886, 4179, 455 

. JM2222, MCJ1171 



Iwiowz^^raw^^/om Alex. MS, B. Arroyos yoriUadecaminos.Comun.Rl 338, ¥3520,^^ 

Wacrostphoma macrosiphon (Torr.) Heller. MD. Vailes y lomerios. Escasa. JM957. 
Mandevilla karwinskii (Muell. Arg.) Hemsi. MS. Laderas. Comiin. MCJII68O, V.^ 


\sdepias hrachystephana Torr. MD, MS. Maleza de areas perturbadas. Escasa. V5607. 

\sdepms linaria Cav. MS, B. Arroyos. Comun. CQ665, HI6174, V4148. 

\sdepias texana Heller. MS, B. Comun. V3895, 5254, VR2200. 

Xsdepias sp. B, MS. Laderas. Escasa. V5254a. 

Zynanchum kunthii (Dene.) Standi. MD. Sobre arbustos bajos. Comun. MCJ 11 7 1 8a, V.^ 

Acourtia nana (Gray) Reveal & Kmg. MR, P. Vailes. Escasa. V4181. 
Acourtia parryi (Gray) Reveal & King. MD. Vailes y lomerios. Escasa. JM2998, V4l63. 
Afoar^i^TOwaWte (D.Don) Turner. MD. Laderas pedregosas. Escasa. JMl 537, 3004, V2887. 
Acourtia ivnghtti (Gray) Reveal & King. MS, B. Vailes y laderas. Comiin. V5350. 
Ageratina callophylla (Blake) King & Rob. MS, MD. Laderas. Comiin. V4l65, 4779. 
Agerattna havanemh (H.B.K.) King & Rob. MS, B. Laderas. Comun. V3092, 3587, 4523, 

Agerattna lunghtit (Gray) King & Rob. MS, B. Laderas. Comiin. V4794, 5440. 
Ageratum. corymhomm Zucc. B. Vailes y laderas. Comiin. V3622, 3948, 4795, 5370. 

Artejutna glauca Pall. B. Arroyos. Escasa. V3063a. 

Artemisia ludovidana Nutt. MS, B. Vailes. Comiin. V3109, 3931, 4160. 
Aster spmosm Benth. MD. Vailes, en represas y arroyos. Escasa. JM303, 781. 

Bahia ahsinthtfolia Benth. var. absinthifolta. MD, P. Vailes. Comiin. JM466, 771, 1034, 

aileya multtradiata Harv. & Gray. MD. Vailes al SW. Comun. V6341. 

arroeteasMigera {Sch 

,aver) Gray. MS, B. Vailes y arroyos. Muy comun. V3100, 3' 

tdens higelovtt Gray. MI 

5, B. Areas perturbadas. Escasa. V3624, 5289. 

idens ferulifolia gacq.) 

DC. MS, B. Arroyos. Escasa. V4487. 

rickellta chloroUpis (Wc 

,ot. & Standi.) Shinners. MS, R Arroyos. Comiin. V3073, 4l( 

ru-kelUa cyltndracea Gr 

ay & Engelm. MS. Arroyos. Escasa. Purp4725, 4728, V465f 

nckellia ladmata Gray 

. MS, B. Arroyos y areas perturbadas. Escasa. V5465. 

. var. conduplicata (Rob.) Turner. MS. Laderas pedregosas. Con 


1167, 5372. Brickeina lemmomt Gray. var. nelsonn (Rob.) Turne 

Escasa. V4805. 


utt. MS. Areas pc 

Erigeron btgelovti Gray. MS 

;. Valles y laderas 

Erigeron chiangii Nesom. 

MS, B. Arroyos, 

Erigeron modestm Gray. M; 

5, B. Lugares hur 

Bnckellia urolepis Blake. B. Comun. V4786, 5290. 

Brkkellia veromcaefolia (H.B.K.) Gray. MS, MD. Escasa. V4166, 4781. 

Calyptocarpus vialis Less. MS, B. Areas perturbadas. Escasa. V5294. 

Centaurea amertcana Nutt. MS. Areas perturbadas. Escasa. V3951, VR2198. 

Chaetopappa bellioides (Gray) Shinners. MD, P. Valles. Comun. V3900, 4725, CQ683, 728. 

Chaetopappa parryi Gray. MS, B. Laderas y paredes rocosas. Comun. V5253, 5302, 5304. 

Chaptalia hololeuca Greenm. MD. Valles y laderas. Kara. V3527. 

Chaptalta texana Greenm. MS, B. Valles. May comun. CQ1057, V3527a, 3977, 5375. 

Chrysactinia mexkana Gray. MD, MS. Valles y laderas. Muy comun. JM617, Rl 336, V5377. 

Chrysactinia pinnata Wats. MS, B. Laderas rocosas. Escasa. V4656, 5234. 

Cirsium undulatum (Nutt.) Spreng. MS. Arroyos y areas perturbadas. Escasa. MCJl l695a, 

Conodinium greggit Gray. MM, MS, P. Areas protegidas. Comun. V3079. 
Conyza coulteri Gray. MS, P. Areas perturbadas. Escasa. V4l42. 
Coreopsis tintoria Nutt. var. ttntoria. B, MS. Areas perturbadas. Escasa. V3544. 
Dichaetophora campestris Gray. MS. Areas perturbadas. Kara. CQ691. 

Dyssodia papposa (Vent.) Hitchc. MS, B, P. Maleza de areas perturbadas. Comun. V4767, 

.adas. Escasa. V3570. 

lun. V3588, 4757. 

as rocosas. Comun. MCJ11700, V4662, 

. Comun CQ735, 1053, V4726, R1327, 


Ertgeron pubescens H.B.K. MS, B, MD, P. Muy comiin. JM4503, CQ734, V4518, 5374. 
FUischmanniapynocephala (Less) King & Rob.. B. Valles y laderas. Comun. V3577, 4780. 
Flourensia cernua DC. MD. Valles. Escasa. JM335. 
Flourensta retinophylla Blake. MS, MD. Valles y laderas. Comun. Purp4728, MCJlOllOa, 

10509a, 11702e, V3555, 4154, 5456. 
Gaillardia gypsophila Turner. MD. Lomerios. Escasa. V5590. 
Gnaphalium roseum H.B.K. B, MS. Laderas y valles. Escasa. VR2239. 
Gnaphalium semiamplexicaule DC. MS, B. Arroyos y areas protegidas. Escasa. V4672, 4754. 
Gnaphalopsis micropoides DC. MD, P, MS. Valles. Comun. JM1018, V2894, 4494, 4742. 
Gnndelia grandtflora Hook. B, MS. Arroyos y areas protegidas. Escasa. C475, V3932, 5291, 

Gochnatia hypoleuca (DC.) Gray. MS, MD, B. Arroyos. Muy comiin. JM767, 2253, V3067. 
Gutterrezia microcephala (DC.) Gray. MD, MS. Areas perturbadas. Comun. V3111, 3899, 

Gutterrezia sarothrae (Pursh.) B. & Rusby. MD. Areas perturbadas. Escasa. C477, V4l57. 
Gutierrezia sphaerocephala Gray. MS, MD, P. Areas perturbadas. Comun. JM141 1, V3915, 

Gymnosperma gluUnosum (Spreng.) Less. MS, MD, B, R Areas perturbadas. Comun JM1070, 

■ra. Comiin. Hinl6555, Purp4708, 

'nium quadridentatum Labil. MD. Areas inundadas. Comiin localmente. JM1022. 
anthella mexicana Gray. B. Areas protegidas. Comun. V3950, 4774, 5275. 
anthus annuus L. MD, MS, R Maleza de areas perturbadas, CI 125 
opsis parvtfolia Gray. MS, B. Valles. Comiin. V3556, 3872, 5365, VR2223. 

ILLARREAL Q., Flora Vascular de la Sierra de la Paila 1 

'eterotheca villosa (Pursh.) Shinners. MS, B, Arroyos y paredes rocosas. Comun. V396 

5300, 5341, VR2221. 
'ymenoxys linearifolia Hook. P, B. Areas percurbadas. Escasa. V4153. 
'ymenoxys scaposa (DC.) Parker, var. scaposa. B, MS, R Valles. Comun. CQ670, MCJ 1 1 68 

V3602,4762, R1326. 
'a ambrostaefolia Gray. Areas perturbadas. Muy comun. V3911, 4691. 
'fea brevtfolta (Gray) Strother. MD. Laderas. Comiin. Hinl6572, JM958, 997, V451 

Lado N. Kara. MCJ 11714a. 
Koanophyllum solidagimfolmm (Gray) King & Rob. MS. Arroyos y laderas rocosas. Escas 

Leucelene ertmdes (Torr.) Greene. MD. Valles y laderas. Escasa. V4133. 

Liatris punctata Hook. var. mexkana Gaiser. B. Rara. V3944. 

Machaeranthera gypsophila Turner. MD, P Valles y lomen'os. Comun. MCJ10123, V415 

Machaeranthera pinnattfida (Hook.) Shinners. var. pinnatifida. MD, R Areas perturbadas. 

JM1936, V3927,6357. 
Melampodium argophyllum (Gray ex Rob.) Blake. MD, MS. Valles. Comun. Purp4730, 

R1479, Hinl6563,V5210. 
Nmlletia edwardsii Gray. MD. Localmente abundante, valles W de la sierra. JM754, V4l 3 

Palafoxia texana DC. var. texana. MD. Areas perturbadas y laderas rocosas. Comun. JM77 

963, 1024, V3597, 3898, 4746, 5212. 
Parthenium argentatum Gray. MD. Laderas bajas. Comun. JM770. V5649. 
Parthenium confertum Gray. var. lyratum (Gray) Roll. MS, B. Areas perturbadas. C78 

Parthenium incanum H.B.K. MD. Valles y laderas bajas. Comun. JM765, V5219. 
Pectis angustifolia Torr. var. tenella (DC.) Keil. MD. Valles. Comiin. R1473, V4138. 
Pinaropappus roseus (Less.) Less. MD, MS. Laderas y valles. Comun. V4631, 6354. 
Porophyllum amplexkaule Engelm. MD, R Laderas rocosas. Escaso. JW083. 
Porophyllumscoparmm Gray. MD. Arroyos. Muy comiin. JM795, V5595. 
Pstlactts tenuis Wats. P, MS. Areas perturbadas. Comiin. V3972, 4139, VR2225. 
Pstiostrophe gnaphalodes DC. MD. Areas perturbadas. Muy comiin. JM796. 
Sanvitalia angustifolia Engelm. B, MS. Areas perturbadas. Comiin. VR2234. 
Sanvitalia ocymioides DC. MS. Areas perturbadas. Comun. V3947, 5323, VR. 
Sartwellia puherula Rydb. MD. Valles y lomen'os. Muy comiin. V5580. 
Sclerocarpusuntsenaits'&^Tvxh. var./r;^/fjfm(Brandg.)Feddma. B, MS. Comiin. V3535, 3910, 

Semcto coahuilensis Greenm. B. Laderas y valles. Rara. V3946, 537L 

iouglasii DC. var. longtlobus (Benth.) Benson. MD. Areas pertubadas, N 

y Wde 

. Comiin. JM340, MCJ10515c, V2895. 

eomexicanus Gray var. neomexicanus . B. Laderas y areas protegidas. Escasa 


veluttna DC. MD. Areas perturbadas. Escasa. V4790. 

oleraceus L. MD, MS. Areas perturbadas, arroyos. Maleza comun. 

tcrantha Lag. B. Areas protegidas y arroyos. Rara. Purp 4721, V5325. 

ata Willd. B, MS. Areas perturbadas. Escasa. V4782. 

losa Lag. MS, B. Valles. Escasa. V3934, 5285, VR2251. 

dmfolta Cav. var. Integra (Blake) Rob. MS, B. Laderas rocosas. Escasa. P 


120 SiDAl6(l) 1994 

Stevia serrata Cav. MS. Areas perturbadas. Comun. V3575. 

Stevia tomentosa H.B.K. B. Areas perturbadas. Comiin. JM1042, V4770. 

Tagetei lucida Cav. B. Areas perturbadas. Escasa. V3874, 5274. 


Theksperma megapotamicum (Spreng.) Kuntze. MD, MS, P. Laderas bajas y valles. Comun 

Thelesperma simplktfolium Gray. MS, P, Areas perturbadas. Comun MCJl 1683a, V3968, 

Thelespetma subaeqi/ale Blake MS, P Areas perturbadas. Comun. V3967, 5248. 
Thymophylla aceiosci (DC ) Strother MD Valles y iomerios. Comun. V4l4l, 5587.6345. 
Thymoph)lL, puiUnhcuh, (DC ) Small var puberula (Rydb.) Strother. MD, MS. Areas 
perturbadas Mu) comun JMI 009, 1017, 1055, 2295, V2893, 3077, 3595, 5313,6346. 

Tnxn ialifouiua Y.z\\ vav cahfonma MD Arroyos y laderas. Comiin. V5 597. 

Ve^besma ihihuahuemn Gray MS, MD Paredes rocosas. Escasa. Purp4694, MCJlOlOSd, 

V3093, 3310 
Yerbenna davmmelMtx^&t MS Laderas Escasa V5287. 
V^rfcj/K^ me//o/^ei (Cav ) Benth & Hook MD, MS Maleza de areas perturbadas. JM1007, 

VerbesmamutopteraUC MS, B Valles y arroyos Comun. V3557, 3966, 4756, 5363. 
Vernoma greggn Gray var ehrenbergti (Gray) Champan &Jones. B. Laderas y orilla de caminos. 

Escasa V3942, 5351 
Vigmera brevifolia Greenm MD Valles y laderas del W de la sierra. Escasa. Endlich9l6 

(MO), Purp4697. 
Viquiera cordifolia Gray. MS. Valles. Escasa. V3079, Purp4693. 

Yiquieradentata{<ZzM)^^^txxg. MD, MS, P. Areas perturbadas. Comun. V3078, 3554, 5339. 
Viquiera greggit (Gray) Blake. MD, MS. Laderas. Comun. MCJI0104, Purp4698, V4152, 

Viquiera longifolia (Rob. & Greenm.) Blake. MS, P. Areas perturbadas. Comun. JM1068, 

2240, V4146, 4776. 
Viquiera stenoloba Blake. MD. Laderas. Muy comun. JM764, V5438, 5596. 
Wedelta acapuicensis Kunth. in H.B.K. var. hispida (Kunth.) Strother. MS. Valles y lomen'os. 

Escasa. V3964. 
Xanthium strumarium L. MD. Maleza de areas perturbadas. Escasa. V4692. 
Xylothamia pseudobaccharis (Blake) Nesom. MD. Valles. Escasa. Hinl6546. 
Zaluzania triloba Gray. MD. Valles y laderas. Comun. JM616, MCJ117l4a. 
Ztnnia acerosa (DC.) Gray. MD. Valles y laderas. Muy comun. JM990, 1038, Hinl6502, 

V5467, 5594. 
Zinnia grandiftora Nuct. MD. Valles y laderas. Comun. V4516, 4639, 5451. 


^ laderas. Escasa. MCJl 1709b. 
eras. Comun. JM1059. 

Cordia bomieri A.DC. MS. Arroyos al E. de la sierra. Kara. V5202. 

Cryptantha mexicana (Brandg.) I.M.Johnst. MD. Arroyos y caminos. Comun. CQ6 

Ueliomptum conferttfoUum (Torr.) Gray. MS, MD. Escasa. V4540. 
Heltotroptum greggii Torr. MS. Arroyos. Escasa. V4126. 

Ueliomptum torreyi I.M.Johnst. MD, MS. Laderas y arroyos. Comun. V4492, 6668. 
Ltthospermum herlandteri I. M. Johnston. MS. Laderas. Comun. MCJ 1 1 694a. 
Lithospermum viride Greene. MS, B. Areas protegidas. Comiin. V5256, 5344. 
Ttquilia canescens (DC.) Rich. MD. Valles y laderas. Muy comun. JM780, V4542. 
Tiqmlia greggii (T. & G.) Rich. MD. Laderas. Comun. JM337, C463. 
Tiqutlia mexicana (Wats) Rich. MD. Valles. Comiin. CQ629, V4543, 4677. 

Brassicaceae (Cruciferae) 

Brasma kaber (DC) Wheeler. MD. Orilia de caminos y arroyos, maleza. Escasa. V5583 

Descuraima pinnata (Walt.) Britt. MD. Areas protegidas. Comun.V6360. 

Eruca sativa Mill. MD, MS. Maleza de areas perturbadas. R796. 

Lepidium virginkum L. MD. Maleza de areas perturbadas. R802. 

Lesquerella argyraea (Gray) Wats. ssp. diffusa (Roll.) Roll. & Shaw. MD. Valles y lade 

pedregosas. Comun. V5 582. 
Lesquerella fendleri (Gray) Wats. MD, MS. Comiin. R799, 1332, V2890. 
Lesquerella purpurea (Gray) Wats, ssp foltosa (Roll.) Roll. MS, B. Areas protegic 

Escasa.V3585, 5373. 
Nerisyrema camporum (Gray) Greene. MD. Valles y laderas. Comiin. CQ790, V2891, 63' 
Nerisyrema linearifolta (Wats.) Greene, var. lineartfolta. MD. Valles y laderas del W y N 

laSierra. Comiin. MCJIOI 11, 10118a, 10510, R1467. 
Sisymbrium auriculatum Gray. MD. Arroyos y areas perturbadas. Escasa. V4129, 6359- 
Sisymbrium irio L. MS, B. Canones y valles con humedad. Escasa. VI 504. 
Sibara vierecki (Schulz.) Roll. MS. Arroyos y laderas con humedad. Escasa.R1332. 

. MD. Areas perturbadas. Muy comiin. R798, V467? 

BurserafagaroidesiH.B.K.) Engler. v^r.fagaroides. MD. Laderas. Escasa. MC 


Ariocarpus fssuratus (Engelm.) Shummann. MD. Rara. 
Astrophytim capricorne (Dietr.) B. & R. MD. Laderas. Escasa. Elizondo294 (. 
Coryphantha borwingii Purp. MD. Laderas y valles. Escasa. JW177, V563 
Coryphantha delaetiana (Quehl) Berger. MD, MS. Valles. Comun. JW153 

V5210a, 5631. 
Coryphantha diffialts (Quehl) Berger.MD, MS. Valles. Comiin. JWl4l, 1: 
Coryphantha echinus (Engelm.) B. & R. MD. Valles. Comiin. Elizondo295 
Coryphantha lam Bremer. MD. Laderas. Escasa. Bremer476-3 (ASU). 
Echmocactus honzonthalonius Lem. MD. Valles y lomerios. Comun. JWl 1 ; 
Echinocereus blamkii (Posel.) Pal. MS. Valles. Escasa. JWI46, 159, 183. 

122 SIDA16(1) 1994 

Echinocereus delaettii Gurke. B. Valles y laderas. Comiin. JW157. 

Echinocereus enneacanthus Engelm. var. duhtm (Engelm.) Benson. MD.MS. Valles y lomerios. 

Comun. JW128. 
Echinocereus enneacanthus Engelm. var. enneacanthus. MD. Lomerios. Comun. JWL66. 
Echinocereus pectinatus (Scheidw.) Engelm. MD. Lomerios y laderas. Comun. JW123. 
Echinocereus stramineus (Engelm.) Engelm. MD. Lomerios. Comun. JWIO6, 116. 
Echinomastus mariposensis Hester. MD. Laderas y valles. Escasa. V5629. 
Epithelantha micromeris (Engelm.) Weber, var. greggit (Engelm.) Ber.MD. Valles y laderas. 

Comun. JW127, 162, V5231a. 
Escobaria dasyacantha (Engelm.) B. & R. MD, MS. Laderas y paredes. Comiin. JW158, 180, 

V5190a, 5269. 
Escobaria tuberculosa (Engelm.) B. & R. MD, MS. Laderas rocosas. Comun. V4798. 
Ferocactus hamatacanthus (Muhl.) B. & R. MD, MS. Laderas y valles. Comun. JW109. 
Ferocactus stainesii (Hook.) B. & R. MD, MS. Laderas. Comiin. JW125. 
Ferocactus uncinatus (Gal.) B. & R. MD. Valles y laderas. Comun. JW156, V5628. 
Grusonia bradtiana Coult. MD. Valles y lomerios. Comiin. V4799. 
Gymnocactus aguirreanus Glass & Foster. MD. Laderas, W de la sierra. Escasa. Glass & Foster 

3044, 2206 (POMA). 
Leuchtenbergia princtpts Hook. MD. Valles y lomerios. Escasa. JW155. 
Lophophora wtliiamsii (Lem.) Coult. MD. Valles. Rara. 
Mammillana chionocephala Purp. MD, MS. Laderas. Escasa. 
Mammillaria melanocentra Posel. var. meiacantha Craig. MD. Valles y laderas. Comiin. 


' imun.JWlOS. 
) B. & R. MD. Laderas y lomerios. Comiin. JW132, V5633. 
Neolloydia smithii (Muehl.) K. & F. MD. Laderas. Comiin. JW131, l65, Eli2ondo478 

Opuntta anteojoensis Pinkava. MD. Valles al N y W de la sierra. Escasa. Lopez sn (ANSM). 

Opuntia grahamii Engelm. MD. Lomerios y laderas bajas. Comiin. V5592. 

Opuntia imhrtcata (Haw.) DC. MD. Valles. Muy comiin. JW120. 

opuntta kleiniae DC. MD.MS. Valles. Muy comiin. JW120. 

Opuntia leptocaulis DC. MD. Valles y lomerios. Muy comiin. JWl 17. 

Opuntia lindheimeri Engelm. var. aciculata (Griff.) Bravo. MS. Laderas. Escasa. 

Opuntia lindheimeri -Engdm. var. lindheimeri. MD, MS. Valles. Comiin. JWIOO, 164, V5634. 

Opuntia lindheimeri Engelm. var. subarmata (Griff) Eliz. & Wehbe. B, MS. Laderas y valles. 

Escasa. JWl 80. 
Opuntia lindheimeri Engelm. var. tricolor (Griff.) Benson. MD, MS. Laderas. Muy comiin. 
Opuntia macrorhiza Engelm. MD. Valles y laderas bajas. Escasa. Eli2ondo468, 469 (ANSM). 
Opuntia microdasys (Lehm.) Pfeiffer. var. microdasys. MD. Laderas y valles. Muy comiin. 

JW102, 148, V5591. 
Opuntia moelleri Berger. MD. Valles al N de la sierra. Comiin. Elizondo342 (ANSM). 
Opuntia phaeacantha Engelm. var dmata (Griff.) Benson & Walk. MD. Valles y laderas. 

Comiin. Espinosa31, 50 (ANSM). 
Opuntia phaeacantha Engelm. var. wz^ji/or Engelm. MD. Valles y laderas. Comiin. Elizondo464 


Ahelia cortacea Hemsl. MS. Canones y lugares protegidos. Kara. MCJ11678. 
Lonkera pilosa WiUd. B. Sobre arbustos. Kara. V4769, VR2210. 


Drymana anomala Wats. B. Lugares humedos. Escasa. V4528. 

Drymaria axillaris Brandg. MD. Arroyos y lomerios. Comiin. V5579. 

Drymaria glandulosa Persl. var. glandulosa. MS, B. Lugares humedos y arroyos. E 

Drymaria polycarpoides Gray. MD. Lomerios. Escasa. S. Vasque24l (ANSM). 
Drymaria subumbellata LM.Johnst. MD, MS. Laderas. Escasa. MCJ 10103, 10515b. 
Paronychia monticola Cory. MD. Laderas y valles. Kara. MCJll681e. 

Schaefferia cuneifolia Gray. MD.MS. Valles y laderas bajas. Escasa. V3104. 

Atriplex canescens (Pursh.) Nutt. MD. Valles y laderas. Escasa. V5622. 
1 disturbio. Escasa. VI 509. 
ishumedasyperturbadas.Comun. V3590,4168, 

Convolvulus eqmtans Benth. MS. Maleza de areas de cultivo. Escasa. V3916, 6367. 
Cuscuta glabnor (Engelm.) Yuncker. MD. Parasita de Vlourensta. Comun. V6337. 
Dichondra argentea H. & B. MD, MS, B. Comun. V3603, 4789, 5358, VR2230. 

Evolvulus alsinoides L. var. hirticulis Torr. MD, MS. Comun. R1468, V3917, 4533, ()(>(i(i. 

Evolvulus sericeus S.Wats. MD. Laderas y arroyos. Escasa. V4497, 5251, 6667. 

ipomoea collina House. MS, B. Sobre arbustos bajos. Comiin. V3923, 4663, 5247, 5347, 

ipomoea costellata Torr. MD. Sobre el suelo o arbustos bajos. Comiin. V45 54, 4690, VR2222. 
ipomoea murtcata (L.) Jacq. MS. Sobre arbustos. Escasa. V. 4552, 4664. 
ipomoea purpurea (L.) Roth. MS. Sobre arbustos de areas perturbadas. Escasa. V4551. 
ipomoea sescossiana Baillon. MS. Arroyos. Escasa. MCJ11690, V4553. 
ipomoea zimmermanit McDonald. MD. Laderas y arroyos. Escasa. Zimmernnanl948 (TEX- 

LL), V3982,6672. 

Echeveria strictiflora Gray. MD. Laderas rocosas. Escasa. H 16 197a, V6669. 
Lenophyllum wembergti Britt. MD, MS. Laderas rocosas. Escasa. MCJlOlOSb, V3072, Al 
Sedum wrightii Gray. MS. Laderas y paredes rocosas de arroyos. Escasa. V4l80, 5369. 
Vtlladia cucullata Rose. MD. Laderas rocosas. Escasa. MCJ10109e, V4l77. 


Forsellesia spmescens (Gray) Greene. MD. Laderas y arroyos. Escasa. H 16 170. 

Skyos an 


L. MS. S 

B.K. MD. Arroyos y areas perturbada 
) Standi. MD. Sobre arbustos bajos o el 
obrearbustos. Rara.V4785. 




MD, MS. Arroyos y canones. Escasa. 


Arbutus xalapensis H.B.K. B. Comun. V3873, 4796. 
Comarostaphylispoltfolia (H.B.K.) Zucc. ssp coahuilensis Hennc 
MCJIOIOI, 11682. 


Acalypha lindheimeri Muell.Arg. B. Arroyos y lugares humedos. Escasa. V3592, VR2228 

Acalypha monostachya Cav. MD, MS, B, P. Comun. CQ783, V4 5 47, 5243. 

Argythamnia humilis (Engelm. & Gray) Muell. Arg. var. humilis. MD, MS. Escasa. V4521 

Bernardia myrcifolia (Scheele) Wats. MD, MS. Canadas. Comun. V3107, 3571, 4678. 
Chtropetalumschiedeanum (Muell.Arg) Pax. MS. Laderas de los arroyos, E de la sierra. Comun 

V3989, 4655. 
Croton diokus Cav. MD, MS. Laderas. Escasa. JM929, 1004. 
Croton frutkulosus Torr. Lugares humedos. Comun. JM985, V3572, 4665, 4741. 
Croton hypoleucus Schltcht. MS, B. Arroyos. Muy comun. MCJ11710, V3108, 3924, 4478 

4788, 5233, VR2199. 
Croton incanus H.B.K. MD, MS. Canones. Muy comiin. V3113, 3594. 
Croton pottsit (Kl.) Muell.Arg. MD. Laderas bajas. Escasa. R1464, VR2220. 
Croton suaveolens Torr. MD, MS, B. Arroyos y laderas. Muy comun. MCJ11713b, 12753 

V3533a, 3926, 5275, VR2211. 
Euphorbia antisyphilitica Zucc. MD. Laderas bajas y valles. Muy comun. V5565, 5627. 
Euphorbia brachycera Engelm. MS, B. Arroyos. Escasa. V5293, 5817, VR2235. 
Euphorbia cinerascens Engelm. MS, MD. Arroyos. Muy comun. CQ786, MCJIOIOO, 11715 

V4515,4751, 5203, 5292. 
Euphorbia cyathophora Murr. B, MS. Lugares protegidos y humedos. V3894, VR2229. 
Euphorbia dentata Michx. MS. Arroyos y areas perturbadas. Comun, V3885, 4527, 4750 

Euphorbia scopulorum Brandg. MD. Laderas bajas del W de la sierra. Comiin. R1478. 
Euphorbia vtllifera Scheele. MS, B. Arroyos. Muy comiin. MCJll681i, V3623, 3897 


,s bajas. Muy comun. JM961. 
'ides Spreng, MD, MS. Arroyos y laderas bajas. MCJ11697, V3922 

4475, VR2236. 
ttllingia treculeanaiUudi. Arg.) l.M.Johnst. MD. Canones. Escasa. JM1392. 
'^ragia amblyodonta (Muell.Arg.) Pax.&Hoffm. MS.MD. Areas perturbadas y arroyos 

Comun. V5296. 

Fabaceae (Leguminosae). Mimosoideae 

Vcacia berlandieri Benth. MS, MD. Arroyos y laderas. Muy comun. C467, V5624. 
Vcacia constricta Gray. MD. OriUa de caminos y laderas bajas. Comiin. C467, JM228(). 
vcacia greggii Gray. MD, MS. Arroyos. Escasa. V5598. 
vcacia hirta T.& G. MS, B. Escasa. V4l34. 

EAL Q., Flora Vascular de la Sierra de la Paila 12 = 

nvermanci Isley. MD. Laderas bajas y arroyos. Comun. V5458. 

^idnla Benth. MS. Valles y arroyos del E de la sierra. Comun. V3884. 

amrutm, Scheele. MD, MS. Arroyos y laderas bajas. Escasa. CQ929, 951, V5577 

^■a umlatj Bcnth MD, MS. Arroyos y laderas. Comun. CQ928, V3881, 4l45 

s. MS. Arroyos y laderas de canones. Comun. CQ909, H1617 

, 3589, 3868. 

h. MS. Valles y arroyos. Escasa. V4536, 5434. 

1. var. emoryanaMXy. Valles y laderas bajas. Comun. V3311, 448 

f^ zygphylla Gray, x M. emoryana Benth. MD. Laderas bajas. Escasa. V5466. 
mglandulosa Torr. var. glandulosa. MD. Arroyos y laderas bajas. Comiin. CQ750. 
2ca media (Marc. & Gal.) Her. MS. Arroyos. Escasa. V5205. 

Fabaceae. Caesalpinoideae 

W^mOTOjmm^Benth.MD. Laderas. Comun.JM334, 992, 1058, H16176,MCJ: 

717, V3610, 4146, 4486, 5461, 6686. 

Ipinia parryi (Fisher) Eifert. MD. Laderas. Escasa.V5589. 

Hum texanum Gray. MD. Arroyos, lado E y N de la sierra. Rara. 

aecrhta greggii (Gray) Heller. MS. Laderas de canones. Comun. V4490, 4695, 

««5e^^z^^/^«f*(Orc.)Eifert.MD.MS. Areas perrurbadas.Comun.MCJ10509,V3584. 

•anseggta oxycarpa Gray. MS. Laderas y arroyos. Escasa. V3975, CQ692. 

bauhiniotdes (Gray) Irwin & Barneby. Valles. Escasa. CQ358, V4522. 

lindheimeriana (Scheele) Irwin & Barneby. MS. Arroyos. Escasa. V3616. 

monozyx (Irwin & Barneby) Irwin & Barneby. MD. Valles NW de la sierra. Escasa. 


Fabaceae. Lotoideae 

lalus sanguineus Rydb. B. Arroyos y areas con disturbio. Rara. V5355. 

mma virgimcum (L.) Benth. B, MS. Sobre arbustos. Escasa. V3990a. 

tnia angustifolta H.B.K. MS, B. Laderas de arroyos. Comun. V4793, 5260, VR2188. 

I caribea Qacq.) Benth. var. edwardsii (Gray) Hassl. MS, B. Lugares hiimedos. Comun. 

586, 3903, 4735, VR2193. 


lea aurea Nutt. MD. Ladera 


, entre arbu 

tos. Escasa. V4772. 

/^^ ^/ra/or H.B.K. var. ^/co/o 

MD. Arroy 

s y canadas. Muy Comu 

V3076, 3541, 



lea frutescens Gray. MD. Lad 


jas. Escasa. 

CQ779^V3958. ^^ 

lea greggii (Gray) H 


lea hospens (Rose) 


MS. Arroyos. Ml 

y Comun. C468, V395 

7, 4170, 5216, 


lea lutea (Cav.) Wi 

d var // 

tea. Iv 

IS. Arroyos 

y laderas bajas. Comun. V4549, 5240. 


lea melantha (Schav 

er) Rydb 


Arroyos. Comun. V4l64, 5463. 


lea nana Torr. var. 


.) Kearn. & 

Peeb. MD. Valles. Escas 

. V6689. 


lea pogonathera Gra 

y. MD. \ 


^ laderas ba 

as. Muy comun. CQ801 

V3534, 3878, 


Dalea radtcans Wats. MS. Arroyo: 
Dalea wrightti Gray. MD. Valles y 

Desmodium grahamit Gray. MS. Arroyos. Escasa. V3962, VR2195. 
Desmodium momexicanum Gray. B, MS. Arroyos y laderas protegidas. Escasa. V5231a. 
Eysenhardtta parvtfolia Brandg. MD. Laderas N y W de la sierra. Comun. MCJ1171i 

Eysenhardtta polystachya (Ore.) Sarg. MS. Arroyos y canadas. Comiin. JM2246, V3930. 
Galactia brachystachya Bench. MD. Sobre arbustos bajos. Comun. CQ680, 744, MCJl 1721 

V4482,4638, VR2189. 
Indigofera acuttfolia Schlecht. MS, B. Arroyos. Comun. CQ854, V3974, 5258. 
Nissolia platycalyx Wats. MS, B. Sobre arbustos. Comun. V3979, 5235, 6687. 
Phaseolus plagiocalyx Harms. B. Enredadera. Escasa. VR2194. 
Phaseolus xanthotrkhus Piper. MS. Laderas, en canones. Escasa. V3961. 
Rhynchosia senna Hook. var. angusttfolta (Gray) Grear. B, MS. Valles. Comun. VR2190. 
Sophora nuttalltana Turner. B. Valles. Comiin. V5824, VR2253. 
Sophora secundiflora (Ort.) Lag. MD, MS, B. Arroyos y laderas. Comun. JM322, HI6I9: 

16597, RI329, MCJ10517, V4724, 5557. 

Quercus grisea Liebm. MS, B. Canones y valles. Comiin. V5825, VR 2209 
Quercm tntricata Trel. MS, MD. Arroyos y caiiones. Comun. JM1048, C464, V3118. 

Quercus invaginata Trel. MS. Arroyos y laderas de canones. Muy comiin. Purp5030, 5029 

JM2336, 3935, C465, V3117, 3563, 4178, 4666, 5238, 53.34, 5570, MCJ11713a. 
Quercus laeta Liebm. B. Valles y arroyos. Rara. V3938, 4784, 5295. 
Quercus laceyt Small. B. Valles canones y laderas. Muy comiin. V3937, 3939, 5356. 
Quercus mohriana Buckl. MS. Canones. Escasa. JM sn. 

Fouquieria splendens Engelm. ssp spkndens. MD. Laderas rocosas y valles. Muy comiin 



Garrya ovata Benth. ssp lindheimeri (Torn) Dahling. B, MS. Caiiones. Escasa. JM1()76, 

MCJ11709a, V3074, 3952, 4792, 6682. 
Garrya wrightit Torr. B. Canones. Escasa. V4791. 

Centaurium calycosum (Buckl.) Fern. var. calycosmn. MS, P, MD. Valles. Comun. HI6I8I. 
C474, V3918, 4159, 5367. 


Namapalmeri Gray. MS. Arroyos y laderas bajas. Comiin. CQ65 1 , 7 16, V45 1 5, 4684, 667 1 . 
Nama undulatum H.B.K. MD, MS. Areas perturbadas. Comiin. CQ627, V6336. 
Phacelia congesta Hook. MS. Arroyos. Escasa. CQ701. 

a-ocarpa Berl. MS, B. . 

VILLARREAL Q., Flora Vascular de la Sierra de la Paila 1 27 


Koeherlinia spmosa Zucc. MD. Arroyos y laderas bajas. Comun. JM319, 1005. 


Kramerta grays Rose & Painter. MD. Laderas bajas W de la sierra. Escasa. HI 6503, V5626. 
Krameria lanceolata Torr. MS, B. Valles. Escasa. V3581, 5832. 
Krameria ramostssima (Gray) Wats. MD. Laderas bajas. Escasa. V4643. 

Lamiaceae (Labiatae) 

Hedeoma costatum Gray var. costatum. B, MS. Paredes rocosas y laderas. Comiin. V3876. 
Hedeoma microphyllum Irving. MS, B. Valles y laderas. Comun. MCjn695, V3875, 5327. 
Hedioma montanum Brandg. MD. Laderas bajas y valles N de la sierra. Comiin. Purp4964, 

MCJIOIU, C461, V4539, 5435. 
Hedeoma nanun (Torr.) Briq. MD. Laderas y valles al W de la sierra. Comiin. V6366. 


V3532, 3920. 
Salvia hallotaeflora Benth. MD, MS. Arroyos. Comiin. V4501, 4608. 
Salvia fannacea Benth. MS. Arroyos. Comiin. Purp4754, V3075, 3529, 4747 
Salvia greggii Gray. B, MS. Laderas y valles. Comiin. MCJll691a, V3888, 53 
Salvia reflexa Hornem. MS. Areas perturbadas y arroyos. Escasa. V3608. 
Salvia regla Cav. MS, B. Laderas. Escasa. V3611, 3887. 
Salvia roemeriana Scheele. B, MS. Arroyos. Comun. V4777, 5334, 5834. 
Salvia tiliaefolia Vahl. B. Areas perturbadas. Escasa. V3547. 
Scutellaria hispidula Rob. MS, MD. Arroyos y laderas. Escasa. MCJ11686. 
ScutellariawrightiiGi^y. MS, B. Valles y laderas bajas, canones. Comiin. V3526, l 

5332, VR2244. 

Linum lewisii Pursh. B. Arroyos. Escasa. JM2306, V3110, 4156. 
Linum nehonii Rose. B. Areas protegidas y arroyos. Escasa. V4739. 
Linum rupestre (Gray) Gray MS, MD. Muy comiin. CQ636, MCJ 10098, lOlC 

4696, 5217, 5249,6349. 
Linum vernale Woot. MD. Laderas bajas y valles. Comiin. V5586. 

Lobelia henricksomi M.C.Johnst. MS. Arroyos. Lugares hiimedos. Escasa. M 


Buddleja marrubifolia Benth. MD. Arroyos y laderas. Comiin. V3883. 
Buddleja scordioides H.B.K. MS. . 
Emorya suaveolens Torr. MS. Arrc 

Spigelia lindheimeri Gray. MS. Arroyos pedregosos. Rara. V4705, 522' 

Cevallia sinuata Lag. MD. Laderas bajas. Escasa. CQ804. 
Eucnide bartonioides Zucc. MD, MS. Laderas de arroyos y paredes rocos 
Eucnide lobata (Hook) Gray MD, MS. Laderas de arroyos y paredes rocc 


Mentzelia asperula Woot. & Standi. MS. Arroyos. Escasa. V5237. 

Mentzelia hispida WiUd. MS, B. Valles y laderas bajas. Comun. V4520. 

Mentzelia mexkana Thomps. & Zavortink. MD. Laderas y valles pedregosos. Escasa. 

Mentzelia saxicola Thomps. & Zavortink. MD. Laderas pedregosas. Escasa. MCJ10105. 

Echinopterys setosa Brandg. MS. Laderas bajas de canones. Comun. Purp4950, MCJ10121, 

Mascagnia cana SmalL MD. Laderas y arroyos. Comun. MCJ10120, V4150, 4524. 
Mascagnia Itlacina (Wats.) Niedenzu in Endl. & Pranh MS, B. Sobre arbustos, paredes 


Abmtlon hypolemim Gray B, MS Escasa V4654, 6679 

Allowtssadula hoh'ieucea (Scheele) Bates MS Areas protegidas Rara. V3869. 

Anoda crenatifolia Ort MS Areas perturbadas Escasa V4660 

Anodapubescem%c\y\itQ\\t MD Arroyos Rara CQ1230 

Anoda thurberi Gt^y MD, MS Areas perturbadas Comun V470 1,4802. 

Batestmalva violacea {Ko5e)Vty^ MD Laderas Escasa V387() 

Hibiscus ioultet I Grs.y MD Laderas rocosas Comun JM760, V4130, 4653, 636L 

Hibiscus martianm Zucc MD, MS Laderas y Valles Comun JM761, V4652. 

Paioma/asioptt^du SLhi^tk MS Areas protegidas Comun V3612, 3913, 4651, 5328. 

Sidahndk'mcN hn^dm cVGray MS Valles \ areas perturbadas Comun. V36l 5, 3959, 5376. 

S ida longipe^ Grjiy MS, B Valles Comun V5215,5319 

Sidaspino^aL MS Areas perturbadas Escasa V^6l5 

SphaeraUea angtiMifnlia (Cav ) D Don MD, B Areas perturbadas Comun. CQ604, 782. 

Sphaerakea endluhii\}\hndit MD Laderas y lomerios Muy comun. EndHch847, V3595, 




a hastulai 

^a Gn 

ly. MD. 

Valles y 



. V4128 




■idea fragran. 

y (Lin 

dl.) Dc 

ne. MD 

. OriUa 

de cam 

inos, are; 

as perturbadas. ( 



n cautifol 

'/.. Sta 


, Onlla 





ff loni^illo, 




All ion la 




-//./ G 

,.,s Mi: 

• Lad era 


. Comu 

n V447 

^ 5224 5452 


• ^)P^<'t,h, 


(Mart .^ 

V Gal ) ^ 

standi ^ 



. Comiin. V357^ 

'omttera angustifolia Torr. MS. Valles y canadas. Comun V4495. 

'raxnw\ cuspidata Torr. B, MS. Canadas y arroyos. Comun. CQ776, 814, R1337, V3954, 

■raxinus greggit Gray. var. greggii. MS. Laderas y arroyos. Muy comun. JM1045, V3953, 

4499, 5299. 
^enodora coulteri Gray. MD. Laderas. Escasa. Purp5020. 
ienodora longifiora Gray. MD. Laderas y valles. Muy comun. JM775, 2274, 223 1 , Hl6 1 73, 

V3880, 4738,VR2215. 

VILLARREAL Q., Flora Vascular de la Sierra de la Paila 129 


Calylophus hartwegn (Benth.) Raven var. hartwegtt. MS, MD. Valles. Comun. MCJ10509b, 

V4733, 5246a, 6339, VR2247. 
Calylophus tubkola (Gray) Raven. MS, MD. Valles y arroyos. Escasa. CQ1051, V3990. 
G^amr^/«Vo/^Raven&Gregory.MS,MD. Arroyos. Comun. CQ1054,V3955, 5255, 5285, 


Orohanche coopen (Gray) Heller. MD. Valles de Larrea y Fouquiena. Escasa. V4513, 6352. 


Oxalts alMcaulis H.B.K. var. pilosa (Nurr.) Eiten. B, MS. Arroyos y areas protegidas. Escasa. 

Oxalts latifolia H.B.K. MS, B. Arroyos. Comun. MCJll681d, V2214. 


Passtflora tenuiloba Engelm. MD. Va 

Argemone fruttcosa Thurb. ex Gray. MD, Valles. Rara. Hin 16582. 
Argemone sanguinea Greene. MD, valles. Escasa. Hin 16600, V636 


Rivina humuiis L. MS, B. Arroyos. Comun. CQ785, V4671, 5227 

Gilia instgms (Brandg.) Cory & Parks. MS, MD. Canadas. Escasa. 
Gilia rigidula Benth. ssp rigidula. \ ' 
Gilta stewardilMJohnst. MD, MS. 

Loeselia coerulea (Cav.) Cav. MD, R ^ 

Poly gala alba Nutt. MD, MS. Canones. 

Polygala barbeyana Chodt. MD, B. Ladei 

4657, 5270, 5639, 6362.VR2246. 
Polygala dolkhocarpa Blake. MD. Arroyos. MCJ10521b. 
Polygala lindheimeri Gray var. lindhelmt 

4749, CQ818,V5827, 6673. 
Polygala macradenia Gray. MD. Lade 

Polygala nudata Brandg. MS, MD. Arrc 

5560, 5615, MCJ10521d, 11720a. 
Polygala parrasana Brandg. MS. Laderas bajas. Escasa. MCJ11699. 
Polygala scoparioides Chodat. MS. Canones y arroyos. Comiin. CQ661, 685, 
Polygala watsonii Chodat. MD, MS. Laderas. Escasa. MCJ11698, V5581. 

Enogonum greggii T. & G. MD. Laderas bajas. Escasa. V3981. 
Eriogonum jamesii Benth. in DC. var. undulatum (Benth.) Stokes ex. Jones. B. \ 

bajas. C 



mun. V3 



Purp4760, JM23 

29, R795 


s. Comur 


703a, V3 



deras ba, 

as. Comun 


Muy comun. JM777, 78 

, CQ659, 

n. Purp4 

762, HI 

168c, V4 


130 SiDA 16(1) 1994 

Port/iLna miiudiihi I.MJohnsc. MD, MS. Valles y arroyos. Muy comiin. V36l4, 3919, 4511, 

Portulaca retina Engelm. MD. Arroyos. Rara. V4510. 

Talinopsis frutescens Gray. MD. Laderas bajas y paredes rocosas Escasa. V4526. 
Talinum aurantiacum Engelm. MD, MS. Valles y laderas bajas. Escasa. V5220. 


Aquilegia longimma Gray. MS, B. Laderas y arroyos. Escasa. V3528. 
Clematis coahinUmn Keil. MS. Arroyos y paredes rocosas. Escasa. V4657. 
Ckmatn drummondii T. & G. MD, MS. Areas perturbadas. Escasa. CQ1055. 
Clematis pitcheri T. & G. Sobre arbustos bajos. Escasa. V3893, VR2226. 
Thalictrum grandtfolium Wats. B, MS. Areas protegidas y hiimedas. Muy comun. CQ905, 
V36ll,3929, 5278. 


Oligomeris linifolia (Vahl.) Macbr. MS. Laderas bajas pedregosas y lugares inundables. 
Comun. JM468, V5600, 6343. 


Ceanothus greggit Gray. MS, B. Valles y laderas. Comun. CQ870, V3609, 5842, VR2227. 

Colubrina greggii Gray. MS, B. Canones y arroyos. Escasa. V3619. 

Condalia encotdes (Gray) M.C.Johnst. MS. Laderas bajas y valles. Comiin. C460, V4l44. 

Condalia warnockit M.C.Johnst. MD. Valles y laderas bajas. Comun 

Condalia viridts LM.Johnst. MS. Canones. Escasa.CQlOOS, V3890, 5841. 

Karwmskiahumboldtiana (Schult.)Zucc. MS, MD. Valles y canadas. Comun. JM993, V3064, 

Rhamnus bettdifolia Greene. B. Canones. Escasa. V4757. 

Rhamnus standleyana C. B. Wolf. B. Valles y canones. Rara. Purp 4968, V5840, yR2207. 
Ziziphus obtusifoUa (T. & G.) Gray. MD.MS. Valles y canones. Comun. JMsn, V5618. 


Amelanchier denticulata (H.B.K.) Koch. MS, B. Laderas bajas, arroyos y canadas. Comun. 

CQ897, V4149, 4771, 5342, 6685. 
Cenocarpus mojadensis Schneid. MS, B. Laderas en canones. Comun.JM2260, 2283, 3272, 

CQ795,MCJ10518, V3081. 
Cercocarpus montanus Raf. var. paucidentatus (Wats.) F.L.Martin. MS, B. Canones. JM2296, 

V3080, 3914,4764, 5782. 
?runus serotina-^hth. var. w>m(Woot & Standi.) Mc.Vaugh. B. Canones. Rara. V3943, 5821. 
Vauquelinia corymbosa H. & B. ssp. heterodon (LM.Johnst.) Hess & Henrick. MS. Canones y 

arroyos. Muy comiin. JMl 5 18, H16171, 16501, MCJlOUOb, 11684, V2888, 4496, 



Bouvardta termfoha (Cav ) Schlecht MS Arroyos. Escasa. V3530, 4476, 5208. 
Coutaportla pailensis ViUarreal MS Paredes rocosas. Rara. V3063, 3305. 
Galium ununiddtum Gray B Arroyos y areas protegidas. Escasa. V4687, 6351. 
Hedyotis aieroui Gray MD Laderas pedregosas y valles. Comiin. JM988, R1335, V3090, 

3536, 3912,4534,4700,4745 
Hedyotis tntritata Fosb MS, MD Laderas pedregosas. Comiin. V3089. 
Redyotis nigricans (Lam.) Fosb. var nigrnans. MS, B. Arroyos. Comiin. V3625, 3904, 4734. 
Hedyotis pa Imeri (Gray) Lewis MS. Arroyos. Escasa. V4546. 

VILLARREAL Q., Flora Vascular de la Sierra de la Paila 

Randia pnnglei Gray. MD, MS. Laderas rocosas y valles. Muy comun.JM9' 

H16178, ¥3106,3307,3569. 
Rdbunium mkrophylhm (Gray) Hems! . MS, B . Arroyos, Comiin. MCJ 11 68 1 h , V 

4645, VR2219. 

Amyns marshii Standi. MS. Canones. Comun.MCJ117l4, V3583, 4766, 6678. 

Choisya palmert Gray. MS. Canones. Escasa. MCJ 1 1691. 

Ptelea tnfoltata L. MS, B. Canones y arroyos. Muy comun. MCJ11702d, R1331, V3531, 

5209, 5568. 
Thamnosma pailensis M.C.Johnst. MD. Laderas bajas, N de la sierra. Kara. MCJ12751, L. 

Woodruf369 (TEX-LL). 
Thamnosma texana (Gray) Torn i. purpurea (Woot. & Standi.) Lundl. MS. Lechos de arroyos. 

Escasa. V4682, 5242. 


Dodonaea viscosa ]a.cq. MS. Lechos de arroyos. Comiin. H 16 169, MCJ 10 119, V3309, 3546, 
Ungnadia specwsa Endl. MS, B. Arroyos y cafiones. Comun. V3580, 5316. 


Bumelia lanuginosa (Michx.) Pers. var. rigida Gray. MS, MD. Canones. Escasa. CQ841, 953, 


Fendlera ngida LM.Johnsc. MS. Laderas rocosas, Escasa. MCJlOlOSc, 11683b, V5226. 
Fendlerella lasiopetala Standi. MS. Laderas rocosas. Escasa. JM1050, MCJ11696, V5818. 


Castela erecta Turp. ssp texana (T. & G.) Cronq. MD. Valles. Kara. JM333, 615, V5619. 
Holacantha stewartii Mull. MD. laderas rocosas. Escasa. JM3897, V6690, 

Casttlleja lanata Gray. MD. Valles y laderas bajas. Escasa. JM321, V6343. 
CasttUeja Itthospermoides H.B.K. MS, MD. Arroyos y laderas pedregosas. Comun. JM3215. 

CQ687, V4755, 5845. 
Hemichaena spinulosa (Wats.) Thieret. MS, MD. Paredes rocosas. Comun. MCJ11679, 

Lamourouxia dasyantha{Ch.z.m. 8cSchi.)^Tn^\:. MD. Laderas pedregosas. Escasa. MCJ 101 l6c, 

Leucophyllum candidum LM.Johnst. MD. Laderas bajas y valles del W de la sierra. Escasa. 

Leucophyllum frutescens (Berl.) LM.Johnst. MS, MD. Valles y laderas bajas. Comiin. V3087, 

Leucophyllum minus Gray. MD. Laderas bajas. Comun. MCJlOlllb, V3086, 4131. 
Maurandya anttrrhimflora H. & B. ssp antirrhiniflora. MD, MS. Sobre arbustos en arroyos. 

Escasa V4632, 5584. 
Maurandya antirrhiniflora H. & B. ssp hedenfolia (Roth.) Elisens. MD. Sobre arbustos de 

valles y canadas. Escasa. JM1081. 
Mecardonia vandelliotdes (H.B.K,) Penn, MD, MS, P, Aroyos y lugares humedos, Escasa, 

Pmstemmi barbatus (Cav.) Roth. MS, B, P Valles y laderas bajas. Comiin. JM2332, V3925, 

Penstemon lanceolatus Benth. MS. Arroyos. Escasa. V4680. 

Penstemon punctatus Brandg. MS, B. Arroyos. Comun. Purp4747, CQ1056, MCJ11685, 

V3601, 3933, 4778, 5236. 
Penstemon triflorus Heller, ssp integrifolms Penn. MS, B. Valles y arroyos. Escasa. R1339. 
Seymeria coahmlana (Penn.) Standi. MS, MD. Valles y laderas bajas. Comun. MCJ10109c, 

Seymeria falcata Turner. y<it.fakata. MS. Valles y laderas bajas. Comun. V3607, 3965, 4685, 

Seymeria virgata (H.B.K.) Benth. MD, MS. Laderas rocosas. Escasa. MCJll681b, VR2205. 

Chamaesaracha comodes (Moric. ex Dunal) Britt. MD. Valles al W de la sierra. Comiin. V5348. 
Chamaesaracha coronopus (Dunal) Gray. MS. Valles y arroyos. Escasa. V4650. 
Chamaesaracha pallida KveKU. MS, MD. Arroyos. Comun. CQ1052, R1334, V4649, 5207, 

6662, VR2249. 
Chamaesaracha sordida (Dunal) Gray. MS, MD. Arroyos. Comun. V3556, 4648. 
Datura tnoxia Mill. MD, MS. Arroyos, orilla de cammos. Comun. JM964, V4694, 5571. 
Datura quercifolta H.B.K. MS, MD. Areas perturbadas y arroyos. Escasa. V4665. 
H««z^;^erw ?ex^«^ (Torr.) Hunziker & Subils. MS. Arroyos. Comun. Rl 340, JM754, V4647, 

Margaranthus solanaceus Schlecht. MS. Areas perturbadas y arroyos. Comun. V4478, 4689, 

Nkotiana glauca Grab. MD. Areas perturbadas y arroyos. Escasa. CQ796. 
Physalis viscosa L. var. cinerascens (Dun.) Waterfall. MS, B. Lechos de arroyos. Comun. V3 5 5 2 . 
Physalis microphysa Gray. B, MS. Canones y valles. Escasa. V5309. 
Quincula lohata (Torr.) Raf. MS. Valles. Comun. V3537. 

Solanum amencanum Mill. MS, MD. Lechos de arroyos. Comun. V3936, 4758. 
Solanum eleagnifolium Cav. MD. Areas perturbadas. Comun. 
Solanum triquetrum Cav. MS, MD. Entre arbustos y lugares protegidos. Comun. V3548, 


Ayenia mkrophylla Gray. MS, MD. Valles y laderas bajas. Escasa. RI459, V5222. 
Ayenia pilosa Cd^toh^l. MS, MD. Valles y laderas bajas. Comun. MCJl 1681, V4477, 4688, 


Aloysta gratissima (Gill. & Hook) Troncoso. MS, MD. Canadas y valles. Comun. CQ798, 

Aloysta wnghtii (Gray) Hellor. MD. Laderas rocosas. Escasa. V4493, VR2203. 
Bouchea spathulata Torr. MS, B, P. Valles. Escasa. MCJ10107, V3099, 6365. 
Citharexylum hrachyanthum (Gray) Gray. MS, MD. Valles. Comun. V3560, 4674. 
Lantana horrida H.B.K. MS. Caiiones y arroyos. Escasa. V4636. 
Lantana macropoda Torr. MD, MS. Valles y laderas bajas. V4635. 
Uppta graveolens H.B.K. MD. Laderas. Muy comiin. V3907, 5197. 
Priva mexkana (L.) Pers. B. Valles y arroyos. Escasa. V5353, VR2224. 
Tetraclea coulteri Gray. MD, MS. Valles y arroyos. Comiin. V5437. 
Verbena bipinnatifida Nutt. MD, MS. Valles. Comiin. 
Verbena elegans H.B.K. MS. Valles. Escasa. V4301. 
Verbena neomexkana (Gray) Small. MS, MD. Arroyos. Escasa. MCJl 168. 


Vkla sororia Willd. B, MS. Arroyos y valles. Areas protegidas. Comiin. R1336, V3985, 4790. 

Phoradendron lanceolatum Engelm. MS, B. Sobre Quercus. Comun. MCJ10102, 11709, 

V3112, 3565, 5303. 
Phoradendron tomentosum (DC.) Gray ssp tomentosum. MS, B. Sobre Acacia. Escasa. V3068. 
Phoradendron viUosum (Nutt.) Nuct. MD, B, MS. Sobre Quercus. Escaso. V3984. 

Cissus incna (Nutt.) Desmoul. MD, MS. Sobre arbustos bajos. Escasa. V4673. 
Parthenoctssus quinquefolia (L.) Planch. MS, B. Paredes rocosas de canones. Kara. V3578. 
Vitis herlandierin^nzh. B,MS. Sobre arbolesyarbusros. Comun. V3525, 3892, 5823, 6661. 


Guaiacum angustifolium Engelm. MD, MS. Valles y canones. Comun. CQ621, V3105. 
Kallstroemia parviflora Morton. MD. Laderas bajas y valles. Comun V5455. 
Larrea tridentata (DC.) Cav. MD. Valles y laderas bajas. Muy comun. JM759. 
Peganum mexkanum Gray. MD. Valles areas perturbadas. Escasa. JM 492. 
Sericodes greggii Gray. MD. Valles y laderas bajas del WS de la sierra. Muy comun. V5602, 

LILIOPSIDA (Monocotideloneas) 

Agave lechuguilla Torr. MD, MS. Laderas rocosas. Muy comiin. JMsn, H16196, V5610. 

Agave macroculmti Todaro. Laderas. Escaso. H16185, Gentry 20050. 

Agave parrasana Berger. MS. Laderas de canones. Escasa. 

Agave scabra Salm-Dyck. ssp scabra. MD, MS. Laderas rocosas. Comiin. V5638 

ve striata Zucc. ssp. falcata (Engelm.) Gentry. MD, MS. Laderas y valles. Comun. 

ndlich879a, V5566, 5612. 

dirion palmen Trel. MD, MS. Laderas rocosas y arroyos. Muy comun. Endlich7. 

eraloe fumfera (Koch.) Trel. MD. Arroyos y valles del E y N de la sierra. Escasa. 

na cespitifera Trel. MD, MS. Laderas bajas rocosas. Comiin. JM2347, V5340. 

a carnerosana (Trel.) McKelvey. MS. Laderas rocosas de canones. Comun. JMs.n., V2897, 

a endlichiana Trel. MD. Valles al W de la sierra. Comiin V2126, 6340. 

dfilifera Chabaud. MD. Valles y arroyos. Comiin. V3986. 
Yucca torreyi Schafer. MD. Laderas pedregosas bajas. Comiin. JM 459- 
Yucca treculeana Carr. MD. Laderas y valles. Comiin. V56l 1. 


Cooperta drummondii Herb. MD. Valles y arroyos. Escasa. CQ781, V6676. 
Zephyranthes longifiora Hemsl. MD. Valles al N. Escasa. V6677. 

Arecaceae (Palmae) 

Hechtta texensis Wats. MD. Laderas y valles rocosos. Muy comiin. V5569, 5613, 5844. 
Tillandsta recurvata L. MD, B. Sobre Fouquieria, Querelas y Pinus. Comiin. CQ817, V31 14, 

Tillandna usneoides (L.) L. B. Sobre Pinm y Quercus. Valles. Comiin. V5837. 


Commelma erecta L. var. angustifolia (Michx.) Fern. MD, MS, B. Laderas rocosas. Comiin. 
V3971, 4488, 4640, 5366. 

134 SiDAl6(l) 1994 

Gibasis linearis (Benth.) Rohw. MD, MS. Valles y laderas pedregosas Comun. V4641, 5338. 
Tradescantia brachyphylla Greenm. MS, MD. Laderas rocosas. Escasa. V5266, VR2237. 
Tradescantia crassifolia Cav. MS. Paredes rocosas de canones. Escasa. V4702, 5337, VR2238. 


Carex schiedeana Kuntz. MS, B. Arroyos y canadas. Comun. MCJ10109a, 1 l691b, 11693, 

V4753, 6674. 
Cyperm acuminatm'X. & H. MS. Lechos de arroyos. Rara. V4500. 


Sisyrinchium scabra C & S. MS, B. Arroyos y canones. Escasa. JM1071, MCJL l681a. 

Alltum kunthii Don. MD. Arroyos. Comun. MCJIOI l6a. 
Echeandia chandleri (Greenm. & Thomps) Cruden. B. Valles y laderas bajas. Comun. V3960, 

Muilla purpusti Brandg. MD. Valles y laderas bajas. Escasa. Purp4959, V3980. 
Schoenocaulon coulteri Baker. MS, B. Valles, canones. Rara. V5245. 
Smilax bona~nox L. MS, B. Sobre arbustos y arboles, canones. Escasa. V3976. 

Dkhromanthus cinnabarinus (Have & Lex . ) Gray. MD. laderas ped regosas . Com lin . MCJ 1 0097 , 

10108c, V4548, 5221, 5279- 
Spiranthes chiangii M.C.Johnst. MS. Laderas rocosas. Rara. MCJlOlOSa. 

Andropogon spadkem Swallen. MS. Valles y arroyos. Escasa. V3084, 3630-6, VR2185. 
Arhtida adscensionis L. MD, P. Valles y laderas. Comun. Ca44, V3627, 4506, 5192. 
Aristida arizonka Vasey. MD. Valles y lomerios. Comun. Ca39, V5385. 
Aristida brawnii Warnock. MD, MS. Valles. Escasa. V5469. 
Artstida purpureanvLZt. var. nealleyi {Vasty) AWrtd. MD, MS. Valles. Comun. V5181, 5470, 

Aristida purpurea Nutt. var. wrightii (Nash) Allred. MD, MS, P. Valles y laderas. Muy comun. 

Aristida ternipes Cav. MD. Valles y laderas. Comun. V4558, 5441. 

Ca56, V5188. 
Bothnochloa lagurotdesiliC.) Hertor. ssp torreyana (Sceud.) Allred & Gould. MD, MS. Comun. 

Bothnochloa saccharoides (Wats) Rydb. ssp reevesti (Gould) Allred & Gould. MS. Valles. Escasa. 

Boutehua barbata Lag. MD, MS. Laderas y lomeios. Muy comiin. Ca9, V5193. 
Bouteloua curtipendula (Michx.) Torr. MD, MS, R Valles y laderas. Comun. Cal2, V5333, 

Bouteloua gracilis (H.B.K.) Lag. MD, R Valles y lomerios. Comun. Ca7, V5352, 5433. 
Bouteloua hirsuta Lag. MS. Valles y arroyos. Comun. Ca63, V5336, 5450. 
Bouteloua johnstonii S2A\Qn. MD. Laderas y valles. Escasa. IMJ8491, 8751, R1483, V4173. 
Bouteloua ramosa Scribn. MD. Laderas y lomerios. Comun. Ca8, V5189. 
Bouteloua trifida Thurb. MD, MS. Valles y arroyos. Comun. Ca6, V3628, 4812, VR2180. 
Brachiaria meziana Hitchc. B, R Valles y areas perturbadas. Escasa. VR2162. 
Brachypodium pringlei Scribn. MS, MD. Laderas. Escasa. MCJ11705, 11705a. 
Bromus anomalus Ropr. MS, B. Valles y laderas. Escasa. Ca51, VR2167. 

BuMoe dactylotdes (Nutt.) Engelm. MD, B, P. Valles y laderas. Comun. VR2163. 

Cenchrus incertus M.A.Curcis. MD, MS. Valles. Escasa. Ca21, VR2154. 

Cenchrus altaris L. MD. Orilla de caminos. Escasa. V5445. 

Cottea pappophoroides Kunth. MS, B. Valles y arroyos. Comun. V5187. 

Cymdon dactylon (L.) Pers. MS, B. Arroyos. Escasa. Ca23. 

Chloris gayana Kunth. MS. Arroyos y areas perturbadas. Escasa. Ca34. 

Chlorh vtrgata Swartz. MD, MS. Areas perturbadas. Escasa. Ca33, V4503. 

Chloris submutka H.B.K. B. Areas perturbadas. Kara. VR2166. 

Dasyochloa pHlchella (H.B.K.) Willd. MD. Valles. Escasa. CalO. 

Dichantheinm oligosanthes (Schult.) Gould var. scriberianum (Nash.) Gould. B.MS. Arroyos y 

canones. Escasa. V3630-7, 5306, VR2181. 
Digitaria californka (Benth.) Herard. MS, B. Valles y arroyos. Comun. Ca37. 
Dzg/>^fw/>/Vc/5;fW;^«(Chase)Stuck. MS, B. Valles y arroyos. Comun. Ca36,V5 191, VR2175. 
Echinocloa colonum (L.) Link. MS. Arroyos. Escasa. Ca36, V3629, VR2169. 
Enneapogon desvauxii Daveau. MS, MD. Valles y laderas. Comun. Ca59, V5231. 
Eragrostis barrelieri Daveau. MD, MS. Valles y arroyos. Comiin. V5194, 5447. 
Eragrostis ciliamnsh Link. MD. Valles y areas perturbadas. Escasa. Ca32. 
Eragrostis curvula (Schard.) Nees. MS. Valles y arroyos. Comun. Ca29. 
Eragrostis intermedta^tt^. MS, MD. Valles. Comun. Ca6l, 27, V3626, 5324, 5383, VR2167. 
Eragrostis mexicana (Hornem.) Link. MS. Arroyos. Escasa. V5470a. 
Eragrostis palmeri Wats. MS. Valles y laderas bajas. Esacasa. V4806. 
Eragrostis pectinacea (Michx.) Nees. MD, MS. Valles y arroyos. Comun. Ca30, V3630-3. 
Erwneuron avenaceum (H.B.K.) Tateoka MD, MS. Valles y laderas. Comun. MCJl 1708a, 

V5196, 5364. 
Erwneuron nealleyi (Vasey) Tateoka. MD, MS. Valles. Comun. Ca3, V5443. VR2155. 
Erwneuron pilosum (Buckl.) Nash. MS. Valles. Escasa. Cal3, VR2179. 
Heteropogon contortus (L.) Beauv. MS, MD. Orilla de caminos. Comun. Ca24, V5195. 

Koeleria pyramidata (Lam.) Beauv. B. Laderas. Kara. Cal5. 

Leptochloa dubta (H.B.K.) Nees. MS, B. Valles y areas perturbadas. Comun. Ca5, V4507, 

5283, 5449. 
Leptoloma cognatum (Schult.) Chase. Arroyos. Comun. Ca28, V5280. 
Lycurus phleoides H.B.K. MD, P Valles y laderas bajas. Comiin. Ca26, V5330. 
Melica montezumae Piper. MS, B. Laderas rocosas. Escasa. MCJl 1704. 
Meto//r^OTex/f^«^(Scribn.)Conert. MS. Valles y arroyos. Comun. Ca58,MCJ10108, 105 14, 

Muhlenbergia dubia Fourn. B. Valles y laderas. Escasa. V5380, VR2184. 
Muhlenbergia emersleyi Vasey. MS, B. Laderas. Escasa. Cal8. 
Muhlenbergia glauca (Nees) Mez. MS, B. Laderas. Escasa. V5284. 
Muhlenbergia lindheimert Hitchc. MS, B. Laderas bajas. Escasa. V5436. 
Muhlenbergia murosperma (DC.) Kunth. MS, Laderas y valles. Comun. Cal6, V5272. 
Muhlenbergia monticola Buckl. MS. Laderas. Comiin. Ca45, MCJl 1705b, 11707, 11708. 
Muhlenbergia porten Scribn. MD. Valles y laderas bajas. Comun. Ca46, V5446. 
Muhlenbergia pubigluma Swallen. MS, B. Laderas. Escasa. V5464. 
Muhlenbergia rigens (Benth.) Hitchc. MS, B. Laderas. Escasa. Cal8. 
Muhlenbergia rigida (H.B.K.) Kunth. MS, B. Valles y laderas. Comiin. Ca47, V5379. 
Muhlenbergia setifolia Vasey. MS, MD. Laderas. Muy comun. Cal9, V4l71, 481 1, 5378, 

Muhlenbergia tenmfolia (H.B.K.) Kunth. iMS, P Valles y laderas. Muy comiin. V3084, 4504, 

Nassella ieuchotrkha (Trin. & Rupr.) Pohl. B, Ms. Valles y laderas bajas. Comun. V526 

Nassella tmutssma (Tnn.) Barkw. B, P. Valles. Escasa. Ca31, V5276. 
?anicum hallii Vasey. MS. Arroyos y areas protegidas. Comiin. Ca43, MCJl 17 l6a, V364( 

4, 5190, VR2164. 
Famcum obtusum H.B.K. MS, P. Valles y lugares inundados. Escaso. Ca57. 
Pappophorum bicolor Fourn. MD. Valles y laderas bajas. Escasa. Ca38, V3630~5, 5431. 
Pappophorum vaginatum Buckl. MD. Valles. Escasa. Ca60. 
Ptptochaetiumfimbrtatum (H.B.K.) Hitchc. B, MS. Comun. Ca52, V5281. 
Poa annua L. MS, B. Lugares hiimedos. Kara. Cal. 
Poa involuta Hitchc. MS, B. Laderas protegidas. Kara. MCJ11706. 

Comun. Ca50, V3987, 4809, VR2158, 2l60. 
Sderopogon brevifolius Phil. MD, P Valles y laderas bajas. Kara. Ca54. 
Setaria grisebachit Fourn. MS. Valles y arroyos. Comun. Ca22, V5184, 5383, VR2159. 
Setaria leucopila (Scribn. & Merr.) Schumann. MS. Arroyos. Comun. Ca20, MCJl 01 2 

11716b, V5182, 5382, VR2182. 
Setaria scheelei (Steud.) Hitchc. B. Valles. Escasa. VR2186. 
Sorghasthrum hrunneum Swallen. MS, B. Laderas. Escasa. V531 1, VR2161. 
Sorghum halepense {\..) Pers. MD. Areas perturbadas. Kara. V3630-2. 
Sporobolus airoides (Torr.) Torr. P, MS, MD. Valles y arroyos. Comun. Ca49, V5183. 
Sporobolus cryptandrus (Torr.) Gray. MS, MD. Valles. Comiin. Ca48, V3631-1, 5246. 
Sporobolus pyramidatus (Lam.) Hitchc. Valles y arroyos. Escasa. Ca42. 
Sporobolus wrightit Munro. Valles. Escasa. 

Stipa aha Swallen. MS. Arroyos. Escasa. MCJ11704a, V5185, 5432, VR2174. 
Stipa eminens Cav. MS, B. Valles y laderas. Comun. Ca55, V5442, VR2172. 
Stipa lobata Swallen. MS. Arroyos. Escasa. VR2168. 
Stipa multinodis Beal. MS, B. Laderas. Comun. MCJl 1707a, V4502. 
Stipa neomexicana (Thurb.) Scribn. MS. Valles y laderas bajas. Kara. Ca62. 
Tragus berteromanus Schult. MS. Areas perturbadas. Escasa. Ca35. 
Tridens muttcus (Torr.) Nash. MS. Valles y arroyos. Comun. Ca4, V4505, 4807, 531 

Trtdens texanus (Wats) Nash. MS, MD. Valles y arroyos. Escasa. Cal4, V3630. 

Agradezco la colaboracion de M. A. Carranza y A. Rodriguez G. por su 
ayuda en las coleccas de campo. A los curadores de los herbarios TEX y MEXU 
por permitirme consultar sus colecciones. La contribucion de J. Valdes R. al 
revisar las gramineas, M. C. Johnston por permitirme revisar sus listados de 
colecta, a B. L. Turner y G. Nesom por sus determinaciones y a T. Wendt por 
la revision detallada 






3RADA. 1991. Las Cacta 

al Autonoma de Mexico. 


.. 1979. Coryphantha lam S 

;p. Nov. A new speoc. 

VILLARREAL Q. , Flora Vascular de la Sierra de la Paila 1 

Cano, G.J. yJ.S.MARROQUlN. 1967. Las gramineasde la Sierra de la Paila, Coah. Bol. S 

Nuev. Hist. Nat. "J. E. Gonzz." 1: 59-106. 
CoRREU, D.S., M.C. Johnston. 1970. The manual of the vascular plants of Texas. Te: 

Research Found. Renner, Texas. 
Crabbh, J.A., A.C. JERMY and J.T. Mickel. 1975. A new generic sequence for t 

pterydophyte herbarium. Fern. Gaz. Il:l4l-l62. 
Cronquist, a. 1981. An integrated system of classification of flowering plants. ColumI 

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Dillon, M. 1976. Systematic study of the genus Flourensia (Asteraceae-Heliantheae). Ph. 

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Fryxell, RA. 1988. Malvaceae of Mexico. Syst. Bot. Monogr. 25:1-522. 
Glass C. and R. Foster. 1972. Gymnocactus aguirreanm a new species from southe 

Coahuila, Mexico. Cact. Succ. J. 44:80-81. 
Gentry, H.S. 1982. Agaves of continental North America. Univ. of Arizona Press. Tucsc 

Henrickson, J. 1981. A new subspecies of Comarostaphylts polifolia (Ericaceae) foi 

Coahuila, Mexico. Madrono 28:33-37. 
Henrickson, J. and D. Flyr. 1 985 . Systematics oiLeucophyllum and Eremogetum (Scrophula 

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38 SiDAl6(l 

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Department of Botany 

University of Georgia 

Athens, GA 30602, U.S. A 

e form of documentation h 

The herbarium at the University of Georgia was founded in the 1920s by 
John N. Reade, and underwent its first reorganization by Joseph H. Pyron 
and Rogers McVaugh in the 1930s. It was expanded under the direction of 
Wilbur H. Duncan who served as curator until his retirement in 1978. 
Samuel B. Jones, Jr., was appointed curator in 1978 and Nancy C. Coile 
served as assistant curator. During 1978-1980, the herbarium was reorga- 
nized and its scope and activities broadened. In addition, the herbarium 
library was formally established. In 1981, Samuel B.Jones was named Direc- 
tor of the herbarium and Nancy C. Coile became the curator. Michael O. 
Moore became curator in 1989, and David E. Giannasi became Director in 
1991. The University of Georgia herbarium serves as a repository of samples 
of southeastern U.S., with emphasis on the flora of the state and has as one 
of Its major goals elucidating the flora of Georgia. During 1979, the her- 
barium staff prepared check-lists for the floras of Clarke County and the state 
of Georgia (Jones and Coile 1979a, 1979b). In 1985, the 2nd edition of the 
Georgia Plant List was issued (Coile and Jones 1 98 5). In 1 985 - 1 986, an elec- 
tronic high density mobile storage system (compactor) was installed in the 
herbarium. This compactor system almost doubled the storage capacity of 
the herbarium and will allow for continued growth of the collection. In 
1988, Jones and Coile published The Distribution of the Vascular Flora of 
Georgia Qones and Coile 1988). This publication has distribution maps by 
SiDA 16(1): 139-151. 1994 

140 SiDAl6(l) 1994 

counties for all native or naturalized plant species occurring in Georgia based 
upon voucher specimens deposited in institutional herbaria. Although 
5,000 copies were printed, demand for this publication has been great, and 
it is now out of print. The publication represented an important next step 
toward the development of a state Flora. 

The herbarium now holds over 206,000 mounted specimens, of which 
162 have been documented as types. This latter number does not include 
topotypes, paratypes, or specimens suspected as being types, but for which 
no literature citation could be found. A number of these specimens are of 
historical significance, particularly collections of 22 taxa made by A.H. 
Curtiss, a prominent early collector in the southeastern United States from 

Catalogued specimens were documented in at least one three ways: (1) 
through citation in the original publication; (2) through citation as a type in 
a secondary publication, and (3) through annotation as a type by a later 
researcher. Where the original or a secondary publication was consulted, the 
location of the holotype is also given. The label data (locality, date, collector, 
collection number), the reference citation, the kind of type, and source(s) of 
documentation are given for each specimen. The following catalogue is 
divided into four sections: Pteridophyta, Gymnospermae, monocotyledons 
and dicotyledons. Within each section, taxa are listed alphabetically by genus. 

Isoetes tagetiformans P.M. Rury, Amer. Fern J. 68:99-108. 1978. isotype. U.S.A. 

Georgia. Columbia C 



rom Columbia Junior 

High School along 

GA 232 and County Rd 2122, 



IS, P.M. Rury &M. 

Tm^^r 259; cited in ( 

mginal publication (HOLOTY 

■pe: NCU). 

Thelypteris pilosi 



raw. var. alabamensis 

Craw., Amer. Fern 

J. 41:15-20. 1951. 

U.S.A. Alabama. 

Winston Co.: in fissures of Pottsviile 

Sandstone on W fork ( 



tble Springs, 26 Nov 1 

9 A9, L.C.Crawford 


iginal publication 



Windham, Contr.Un 

iv. Michigan Herb. 

19:59. 1993. isotype. 


A. Quebec. Gaspe 

Co.: on sandstone sea 

-cliffs, 19-21 Aug 


dd 25351: 

ype by M.D. Windha, 

m, 1993. 


Podocarpus lucie 

nii Laube 


!0. I960. ISOTYPE. NE 


scattered and locally 


on slopes above ar 

id N of Riviere Bleuc 

'. River, 5 mi from 

confluence with Yat< 

, River, 1 

Oct 1957, D.J 

. de Laubenfels P737, 

: cited in original 

:ation (holotype: GH). 



:tris lutea Small forma albiflora E.T. Browne, Rhodora 63:305. 
. Georgia. Chatham Co.: low ground at edge of Cypress stand 15 

Moore and Giannasi, Type specimens, University of Georgia Herbarium 14 1 

Savannah City Hall, NW of Old Ogeechee Canal on Bamboo Farm-Pembroke Road, 1 3 Jun 
1958, W.H. Duncan 20982; cited in original publication. 

Allium passeyi N.H. Holmgren & A.H. Holmgren, Brittonia 26:309- 1974. isotype. 
U.S.A. Utah. Box Elder Co.: SW part ofHowellValley,TllN,R5W,S7,13Jun I960, A.R 
Holmgren, H.B. Passey & VK. Hugie 13123, cited in original publication (holotype: NY). 

Andropogon brachystachyus Chapm., Fl. S U.S. ed. 2:688. 1 883. isolectotype. U.S.A. 
Florida; dry pine barrens near Jacksonville, Oct no year, A.H. Curtiss 3632; annotated as 
isolectotype by C.S. Campbell, 1980. 

Andropogon virginicus L. var. glaucus Hack, in DC, Monogr. Phan. 6:411. 1889. 
ISOTYPE. U.S.A. Florida: dry pine barrens near Jacksonville, Nov no year, A.H. Curtiss 
3638h\ annotated as isotype by C.S. Campbell, 1980. 

Aristida lanuginosa Scribner ex Hitchc, Contr. U.S. Natl. Herb. 17:278. 1913. 
ISOTYPE. MEXICO. Jalisco: hills near Guadalajara, 28 Oct 1889, C.G. Prmgle2375; cited 

[. S. Calif Acad. Sci. 37:1. 1938. isotype. U.S.A. 
Charleston Park, 2 1 Jul 1937, /, W. Clokey 7479; 
cited in original publication. 

Carex X absconditiformis Fernald, Rhodora 44:387. 1942. isotype. U.S.A. Virginia. 
Sussex Co.: bottomland woe 

1980 (holotype: GH). 

Carex amplisquama F.J. ] 

original publication (holotype: US). 

Carex austrocaroliniana EM. Bailey, Am. J. Sci. 45:173. 1843. isotype. U.S.A. 

Bryson, 1980. 

Carex digitalis Willd. var. asymetrica Fernald, Rhodora 43:544. 1941. isotype. U.S.A. 

NW of Applewhite Church, 8 May 1940, M.L. Fernald andB. Long 11791; cited in original 
publication, annotated as isotype by C.T. Bryson, 1980 (holotype: GH). 

Carex interior EM. Bailey var. charlestonensis Clokey, Bull. S. Calif Acad. Sci. 38:1. 
1939. isotype. U.S.A. Nevada. Clark Co.: Charleston Mountains, Charleston Peak, damp 
soil by spring, 19 Jun 1937, /. W. Clokey 7468; annotated as isotype by A.A. Reznicek, 1978. 

Carex lucorum Willd. ex Link var. austrolucorum J.H. Rettig, Sida 13:449-450. 
1 989. holotype. U.S.A. North Carolina. Buncombe Co.: along NC 9, 5.0 mi N of jet. with 
US 74 in town of Bat Cave, 6 May 1986,7.R Retiig 1534; cited in original publication, 
annotated as holotype by J.H. Rettig, 1988. 

Carex manhartii Bryson, Castanea 50:15-17. 1985. holotype & isotype. U.S.A. North 
Carolina. Macon Co.: CoweetaHydrologic Lab, Stewart Trail, 17 May 1981, J.i?. Manhart 
293; annotated as holotype and isotype by C.T. Bryson, 1985. 

Carex mohriana Mackenzie, N. Amer. Fl. 18:106. 1931. isotype. U.S.A. Florida: 
swamp near Wauchula, 15 Apr 1901, A.H. Curtiss 6761; annotated as isotype by A.A. 
Reznicek, 1978. 

Carex shortiana Dewey, Mss. Dewey, Car. XXX, p. 60. isotype. U.S.A. Kentucky: 
Lexington, no date, Short s.n.; annotated as isotype by S. McDaniel, 1971. 

Cyperus granitophilus McVaugh, Castanea 2:103. 1937. holotype. U.S.A. Georgia. 
Walton Co.: sandy depression in granite rocks, 1 mi E of Loganville, 3 Oct 1936,J.R Pyron 
& R. McVaugh 977; cited in original publication. 


anta Cruz, Chiquitos, Est. San Ignacio, 


,f San Jose, 17^ 

W, 1 Feb IS 

186,7: Kiheen i72S; cited: 

in original! 



ium umbilicatum C Pari 

1 ssp. mom 

)Stolum C. Pa 

Gap, 6 May 

\96\J.W. Hardin 2360- i 

;ited in ori^ 

;inal public 

;ation (holoty 


ium umbilicatum C. Pari 

licatum, Brit 

ell Co.: along slope abov( 

W of States 

villeoffl40, 13 May 196: 

i J. W.Han 

:ited in origin 

(holotype: NCSC). 


ylis diphylla (Retz.) Vahl 

ssp. diffus 

a D.B. Wa 


FL 121, Wi 

orner of Paynes Prairie, 1 

3 Aug 196 

6, D.B. Ward 5867; cit. 


(holotype: FLAS). 


ia blephariglottis (Willd. 

. inteerilaj 

bia Correll, B< 


m sphagnum V 


Zhitley City, 27 Aug 1940 

,P.T. McFarland&H.J 

'. Rogers 97; ci, 


nesii M.G. Chung, Ann 

. Missouri 

Bot. Card 

. 76:920. m 

KOREA: Kyeongsan Nam Do Provin 

ice: Namhae Gun, San. 

gu Myeon, Mt 

^alm Creek near Cape Romano, Jul no year, A.H. C//»Ym 2705; annotated as isotype by R.R. 
rlaynes, 1974. 

Nolina atopocarpa Bartlett, Rhodora 11:81. 1909. isotype. U.S.A. Florida: Indian 
liver, Eau Gallie, no date, A.H. Curttss 3 702; cited in original publication (holotype: GH). 

Rhynchosporacurtissii Britton ex Small, Fl. SE U.S. 195, 1327. 1903. isotype. U.S.A. 
Florida: low roadside near Milton, 8 Jul 1897, A.H. Curtiss 3929; annotated as isotype by 
i.M. Kooyman, 1980. 

Rhynchospora oligantha A. Gray var. breviseta Gale, Rhodora 46:128-130, 1944. 
sotype. U.S.A. Florida: damp pine barrens near Jacksonville, 19 Jun 1896, A.H. Curtns 
)687; cited in original publication, annotated as isotype by S.M. Kooyman, 1980 (holo- 
type: GH). 

n even at low stage, Little River above AL 35 bridge, 27 Jul 1977, R. Krai 60639; cited in 
>riginal publication (holotype: US). 
Scirpus rubiginosus Beetle, Amer. J. Bot. 28:697. 1941. isotype. U.S.A. California. 

)lius R.M. Harper, 

sJ.D. Freeman, Brittonia 27: 17-18. 1975 

Moore and Giannasi, Type specimens, University of Georgia herbarium 143 

Trillium foetidissimum J.D. Freeman, Bnttonia 27:31-32. 1975. jsotype. U.S.A. 
Mississippi. Adams Co.: in loess ravines under rich hardwoods S of US 61-84-98, 1.2 mi E 
of Sc. Catherine Creek in Natchez, 30 Mar 1967, J.D. Freeman 535\ cited in original 
publication, annotated as isotype by J.D. Freeman, 1968 (holotype: GH). 

Trillium lancifolium Raf., Autik. bot.: 132. 1840. isoneotype. U.S.A. Florida. Gadsden 
Co.: moist slopes in woods, Chattahoochee, 14 Mar 1901, A.H. Curtiss 6747; cited as 
isoneotype by J.D. Freeman, 1975, Brittoma 27:10 (neotype: GH). 

Trillium persistans W. Duncan, Rhodora 73:244-246. 1971. ISOTYPE. U.S.A. Georgia. 
Rabun Co.: open deciduous woods on S facing slope of Tallulah Gorge, 23 Mar 1971, W.H. 
Duncan 23548; cited in original publication (holotype: US). 

Trillium reliquum J.D. Freeman, Brittonia 27:21. 1975. isotype. U.S.A. Georgia. 
Columbia Co: (Richmond Co. on label and original publication, but Freeman says 
Columbia), rich woods on bluff slope at mouth of ravine, E facing bluff of Savannah River, 
ca. 5 mi above Augusta, 30 Mar 1968, /D. Freeman (j25; cited in original publication, 
annotated as isotype by J.D. Freeman, 1967 (holotype: GH). 

Yucca smalliana Fernald, Rhodora 46:8. 1 944. isotype. U.S.A. Florida: sandy soil near 
Jacksonville, May no year, A.H. Curtiss 2930, cited in original publication (holotype: GH). 


Agalinis acuta Penneli, Bull. Torrey Bot. Club 42:338. 1915. isotype. U.S.A. Massa- 
chusetts. Martha's Vineyard, Edgartown, dry sandy downs, 12 Sep 1901, M.L. Fernald 43; 
cited in original publication, annotated as isotype by J.M. Canne, 1985 (holotype: US). 

Agalinis edwardsiana Penneli, Proc. Acad. Nat. Sci. Philadelphia 73:522. 1922. 
isotype. U.S.A. Texas. Kendall Co.: dry adobe hills 5 mi NE of Boerne, 16 Sep 1913, /^ W: 
Penneli 5482; cited in original publication (holotype: PENN). 

Agalinis homolantha Penneli, Proc. Acad. Nat. Sci. Philadelphia 73:525. 1922. 
ISOTYPE. U.S.A. Texas. Colorado Co.: sandy oak woodland NW of Sheridan along SA & AP 
railroad, 21 Sep 1913, F.W. Penneli 5522; cited in original publication (holotype: PENN). 

Amorpha georgiana Wilbur, Rhodora 56:261-263. 1954. isotype. U.S.A. Georgia. 
Telfair Co.: 3 mi NW of Lumber City on US 23, dry savannah and river bank, 9 May 1953, 
R.L. Wilbur 3158; cited in original publication (holotype: GH). 

Amorpha ouachitensis Wilbur, Rhodora 77:394-397. 1975. isotype. U.S.A. Okla- 
homa. LeFlore Co. : dry rocky (sandstone) hills. Page, 23 Sep 1 92 1 , E.J. Palmer 205 72; cited 
in original publication, annotated as isotype by R.L. Wilbur, 1973 (holotype: A). 

Amorpha virgata Small, Bull. Torrey Bot. Club 21:17. 1894. isolectotype. U.S.A. 
Georgia. DeKalb Co.: NW slope of Stone Mountain, 3 Jul 1893,/^ Small s.n.; cited as 
isolectotype by R.L. Wilbur, 1977, Rhodora 77:398 (lectotype: NY). 

Aquilegia scopulorum Tidestrom ssp. perplexans Clokey, Bull. S. Calif. Acad. Sci. 
37:3. 1939. isotype. U.S.A. Nevada. Clark Co.: Charleston Mountains, Charleston Peak, 
broken rocky slope above timberline, 16 Jul 1936, l.W. Clokey 7094; cited in original 

Aster erectus E.S. Burgess, Mem. Torrey Bot. Club 13:147. 1906. isolectotype. U.S.A. 
New York: vicinity of New York City, Bryn Mawr Park, 26 Sep 1 896, E.S. Burgess s. n. ; cited 
as isolectotype by W.F. Lamboy & A.G. Jones, 1987, Brittonia 39:290. 

Aster jonesiae Lamboy, Syst. Bot. 13:192-193. 1988. holotype & isotype. U.S.A. 
Georgia. Upson Co.: oak-hickory woods at E base of Pine Mountain, near Pasley Shoals, W 
ofThomaston, 10 Sep 1947, A. CroK^/^^ij/ 4694; cited in original publication, annotated as 
holotype and isotype by W.F. Lamboy, 1988. 

Aster roscidus E.S. Burgess in Britton & Brown, 111. El. N U.S. 3:360. 1898. 
isolectotype. U.S.A. New York: Bryn Mawr Park, 14 Sep 1897 (1898?), £.5. Burgess s.n.\ 
cited as isolectotype by W.F. Lamboy & A.G. Jones, 1987, Brittonia 39:290. 

Aster tenuifolius L. var. aphyllus R. Long, Rhodora 72:40-41. 1970. isotype. U.S.A. 
Florida. Hillsborough Co. : NW of Tampa, S of FL 580 and W of Rocky Creek, 24 Dec 1962, 
0. Lakela 256lO\ cited in original publication, annotated as isotype by S. Sundberg, 1986 

Astragalus aequalis Clokey, Madrono 6:215. 1942. isotype. U.S.A. Nevada. Clark Co.: 
Charleston Mountains, hillside Juniper belt, Harris Springs Road, 4 Jun 1937, /. W. Clokey 
7372; cited in original publication. 

Astragalus hemigyrus Clokey, Madrono 6:220. 1942. isotype. U.S.A. Nevada. Clark 
Co.: Charleston Mountains, rock ledges, Larrea belt, S of Indian Springs, 18 Apr 1939, /• W. 
Clokey 8409; cited in original publication. 

Baptisia arachnifera W. Duncan, Rhodora 46:29-31. 1944. isotype. U.S.A. Georgia. 
Wayne Co.: sandy soil in open pine woods 10 mi SofJesup,4jul 1943, W:H.D««f^K ^693; 
cited in original publication (holotype: GH). 

Baptisia pendula Larisey, Ann. Missouri Bot. Gard. 27:171. 1940. isotype. U.S.A. 
Georgia. Decatur Co.: dry open woods bordering the Flint River near Bainbridge, 19 Jul 
1901, A.H. Curtiss 6810; annotated as isotype by D. Isely, 1978. 

Bidens mariana S.F. Blake, Rhodora 31:88. 1929. isotype. U.S.A. Maryland: sandy 
shore of Northeast River near Carpenters Point, Charleston, 17 Sep 1926, S.F. Blake 9698; 
cited in original publication (holotype: US). 

Bigelowia nuttallii L.C. Anderson, Sida 3:460-461. 1970. isotype. U.S.A. Georgia. 
Tattnall Co.: W bank of Ohoopee River 3.5 mi WofReidsviUe, 12 Nov 1969, L.C Anderson 
3455; cited in original publication (holotype: KSC). 

Bumelia thornei Cronquist, Castanea 14:103. 1949. holotype. U.S.A. Georgia. Early 
Co.: in dry live oak woods by cypress swamp, 1 mi E of Nantz Spring, 22 Oct 1947, R.F. 
Thorne 7345; cited in original publication. 

Calycanthus brockiana Ferry & Ferry f , Sida 12:339-1987. holotype. U.S.A. Georgia. 
Lumpkin Co.: 12.8 mi SW of Dahlonega, S of E-W dirt farm road in open deciduous forest, 
10 May 1982, R.J. Perry Sr. 637, annotated as holotype by R.J. Ferry & R.J. Ferry f, 1987. 

CastillejachristiiN.H. Holmgren, Bull. Torrey Bot. Club 100:91. 1973. isotype. U.S.A. 
Idaho. Cassia Co.: Cache Peak Range, Harrison Mountain, near top, T13S, R24E, S9, 12 
Jul 1966, N.H. Holmgren &J.L. Reveal 2866; cited in original publication (holotype: NY). 

Castilleja clokeyi Pennell, Proc. Acad. Nat. Sci. Philadelphia 89:420. 1938. isotype. 
U.S.A. Nevada. Clark Co.: Kyle Canyon, gravelly loam on N slope, 8 Jul 1936, /. W. Clokey 
& C.B. Clokey 7322; cited in original publication. 

Cirsium clokeyi S.F. Blake, Proc. Biol. Soc. Wash. 49:8. 1938. isotype. U.S.A. Nevada. 
Clark Co.: Charleston Mountains, ridge near peak, gravelly open slopes, 6 Aug 1937, /. W. 
Clokey 7456; cited in original publication (holotype: US). 

Clematis morefieldii Krai, Ann. Missouri Bot. Gard. 74:665. 1987. isotype. U.S.A. 

Moore and Giannasi, Type specimens, University of Georgia 
of Deborah Ave., 0.75 mi SSW of Red Top Mountam, 17 Jun 1^ 

Clitoria fragrans Small, Torreya 26:57. 1926. isotype?. U.S.A. Florida. Highlands Co.: 
sandhills near DeSoco City, 20 May 1925, J.K. S?nall & E.T. Wherry 12626; note on label 
states that Gleason could not find the type at NY in 1945, annotated as possible isotype by 

Cordylanthus glandulosus Pennell & Clokey, Proc. Acad. Nat. Sci. Philadelphia 90: 11. 
1938. ISOTYPE. U.S.A.Nevada. Clark Co.: Charleston Mountains, brushy hills,JuniperBelt, 
Harris Springs Road, 16 Jul 1937, 1.W. Clokey 7713; cited in original publication. 

Coreopsis leavenworthii T. & G. var. curtissii Sherff, Bot. Gaz, 94:592. 1933. isotype. 
U.S.A. Florida: dry pasture ground along the Suwannee River near Stanford, 23 Oct 1900, 
A.H. Curtiss 6734; annotated as isotype by E.B. Smith, 1976. 

Coreopsis longifoliaSmall var. godfreyi Sherff, Bot. Leafl. 6:3. 1952. HOLOTYPE. U.S.A. 
Georgia. Wheeler Co.: boggy, sphagnous depression in sandhills 7 mi S of McRae, 14 Oct 
1950, R.K. Godfrey 50793; annotated as "type" by E.E. Sherff, no date. 

Coreopsis saxicola Alex. var. duncanii Sherff, Bot. Leafl. 6:2. 1952. holotype. U.S.A. 
Georgia. Oglethorpe Co.: shallow soil at edge of thickly vegetated drain at Echols Mill, SE 
of Point Peter, 4 Aug l94l,W.H. Duncan 3832; annotated as "type" by E.E. Sherff, no date. 

T.B. Croat 19382; cited in original publication (holotype: NY). 

Dedeckera eurekensis Rev. & J.T Howell, Brittonia 28:246-248. 1976. isotype. 

Valley sand dunes and 3.5 air mi NW of Marble Valley VABM 7559 in TIOS, R40E, 29 Jul 
1975, y.L, Reveal, M.C. DeDecker & RW. DeDecker 3909; cited in original publication 
(holotype: US). 

Dicerandra frutescens Shinners, Sida 1:89-90. 1962. isotype. U.S.A. Florida. High- 
lands Co.: 20 mi S of Sebring along Route 27, 28 Aug 1953, F.H. Sargent 6600; cited in 
original publication, annotated as isotype by R.B. Huck, 1984 (holotype: SMU). 

Dipsacus sylvestris Hudson f. albidus Steyerm., Rhodora 60:175. 1958. iscjtype. 

celiopsis nudicaulis (A. Gray) Nelson var. corrugata Cronquist, Bull. Torrey Bot. 
99:246. 197 3. ISOTYPE. U.S.A. Nevada. Nye Co.: about outcrops ofpale hard limestone 
rrea desert about 1 5 (airline) mi SE of Lathrop Wells, T17S, R5 IE, about S20, 20 Apr 
, A. Cronquist 10648; cited in original publication (holotype: NY). 
logonum aliquantum Rev., Phytologia 34:460-461. 1976. isotype. U.S.A. New 
:o. Colfax Co.: 0.3 mi W of NM 21, 4.4 mi S of US 64 at Cimmaron on the PhUmont 
Ranch, 14 Jul 1972,7.L. Reveal 277; cited in original publication (holotype: US). 
ogonum capistratum Rev., Phytologia 66:254. 1989- isotype. U.S.A. Idaho. Custer 
Antelope Pass, upper end of Cooper Basin on a ridge SW of the pass, 1 1 Jul 1975,J.L. 

ogonum lewisii Rev., Great Basin Nat. 45:277. 1985. isotype. U.S.A. Nevada. Elko 
Vhite Elephant Butte, S of Elk Mountain, S4, T46N, R16E, 30 Jul \9l6j.L. Reveal 

146 SiDA 16(1) 1994 

Eriogonum ochrocephalum S. Watson van alexanderae Rev., Great Basin Naturalist 
45:276. 1985. isotype. U.S.A. Nevada. Lyon Co.: along NV 3 in Wilson Canyon, between 
Smith and Mason, 12.8 mi NE of Smith, 21 Jun 1978, J. L. Repeal 4737; cited in original 

Eriogonum ovalifolium Nutt. var. pansum Rev., Phytologia 66:259. 1989- isotype. 
U.S.A. Idaho. Boise Co.: along ID 21 ca. 12.5 mi S of Lowman, near West Ford Creek at 
milepost 59.5, 12 Jul 1975,7,L, Reveal & B.J. Ertter 3883; cited in original publication. 

Euphorbia graciliorCronquist.Castanea 14:102-103. 1949- isotype. U.S.A. Georgia. 
Tattnall Co.: sandhills 3 mi NW of Reidsville, 16 Jun 1948, A. Cronqimt 5334; cited in 
original publication, annotated as isotype by M.J. Huft, 1979 (holotype: US). 

Flaveria linearis Lag. var. 





.rts 39:289. 1903. 

SYNTYPE. U.S.A. Florida. Pa 

Im Beach Co.: s 


ofLake Worth near Pal 

mBeach, 31 Aug 

189^,A.H. Curuss 5524; ck 

ed as type 



d publication, annotate 

das type by A.M 

Powell, no date. 

Forestiera pubescens ^ 

Jutt. var. 



ia Shinners, Field & L 

ab. 18:99. 1950. 

isotype. U.S.A. Texas. Bosc 

5ue Co.: t 

ver limestone 12 5 mi 

NNE of Walnut 

Springs, 26 May 1949, L.H. 



rotated as isotype by C. 

J.Brooks, 1975. 

Galacda minor W. Dune 

:an, Phytc 


U.S.A. Georgia. 

Long Co.: sandhills adjacent 





Dum-an 16993; eked inorigii 

lal public 



stated as holotype by W. 

H.Duncan, 1977. 


i, original 

.South Carolina. 

Williamsburg Co.: floodplai 

.nk of 

the 5 

kntee River along SC 17, 0.6 mi N of the 

Berkeley Co. line, about 2.7 mi N of Jamestown, 6 Apr 1975 J. L. Reveal & M.J. Seldtn3832; 
cited by J. Reveal and M. Seldin, 1975, Taxon 25:135. 

Helenium floridanum Fernald, Rhodora 45:494. 1943. isotype. U.S.A. Florida. 
Hernando Co.: low open woods near Fitzgerald, l4 Jun 1900, A.H. Curtiss 6663; cited in 
original publication, annotated as isotype by W.M. Bierner, 1971 (holotype: GH). 

Hexastylis shuttleworthii (Britton & Baker) Small var. harperi Gaddy, Sida 12:54. 
1987. isotype. U.S.A. Georcsia. Madison Co.: acid bog just N of GA 106, l4.2 mi NE of 
Athens, 9 May 1986, L.L. Caddy j.«,; cited in original publicarion (holotype: CLEMS). 

Hoita hirsuta Rusby, Mem. New York Bot. Gard. 7: 261. 1927 isotype. BOLIVIA: 
Pongo, 12 Jul 1921, O.E. White 154; annotated as isotype by J. Grimes, 1990. 

Hymenoxys turned K.R Parker, Phytologia 20:192. 1970. isotype. U.S.A. Texas. 
Karnes Co.: limestone soil along roadside 2 mi N of Karnes City, 21 Apr 1965, 6,L, Turmr 
5154; cited in original publication (holotype: US). 

'[]un I960, R.K.Godfrey 


38:372. 1936. isotype. 

Hypericum adpressum Barton var. spon 

giosum I 

isotype. U.S.A. Massachusetts. Barnstable C< 

..: sandy r 

1901, G,G. Kennedy, E.F. Williams & ML. 

Hypericum canadense L. var. galiiform 

e Fernald 

type. U.S.A. Virginia. Sussex Co.: sandy ai 

id peaty 

■r B.Long. 

Hypericum exile P. Adams, Contr. Gray Herb. 189 

Gulf Co.: 2.5 mi SE of Port St. Joe, sandy dry 

soil of pin 

456; cited in original publication (holotype 


Hypericum lissophloeus P Adams, Con, 


Florida. Bay Co.: shores of Merial Lake, 10 m 


&J.N. Triplett 59844; cited in original publ 

ication (H 

Hypericum mutilum L. var. latisepalui 

m Female 

J.S.A. Florida. Duvall Co.: low grounds, Jun-Aug no year, A.H. Ciirtm 264*, cited in 

Ilex decidua Walter var. curtissii Fernald, Bot. Gaz. 33:155. 1902. isotype. U.S.A. 
LORiDA. Suwannee Co.: woods along the Suwannee River near Branford, 24 Oct 1900, A.H. 
'miss 6736; annotated as isotype by R.C. Clark, 1983. 

Ilex glabra (L.) A. Gray f. leucocarpa F.W. Woods, Rhodora 58:25-26. 1956. isotype. 
F.S.A. Florida. Jackson Co.: open woodland 4 mi S of Marianna, 20 Jan 19^5, F.W. Woods 

Justiciacooleyi E.C. 

LORIDA. Hernando Co.: 
48/; cited in original I 

Lesquerellastonensis Rollins, Rhodora 57:255-256. 1955. isotype. U.S.A. Tennessee. 
Lutherford Co.: field near E fork of Stones River, Walterhill, 26 Apr 1955, R.C. Rollins 
5176; cited in original publication (holotype: GH). 

Lesquerella thamnophila Rollins & Shaw, The genus Lesquerella:?>6 . 1973. isotype. 
J.S.A. Texas. Zapata Co.: sandy soil 3 mi SE of Zapata, 19 Apr 1959, R.C. Rollins &D.S. 
'orrell 3949; cited in original publication. 

Lobelia harrisii Urban, Symb. Ant. 5:520. 1908. syntype. Jamaica: near Troy, 28 Jun 

Ludwigia ravenii C.I. Peng, Syst. Bot. 9:129. 1984. isotype. U.S.A. South Carolina. 
erkeley Co. : roadside sedgy, sandy ditch, 0. 1-0.2 mi SW of the jet. of County Road 1 6 and 
ounty Road 6, on County Road 16 W ofMoncks Corner, 9 Sep 1982, C.l. Peng 4402; cited 


anus Sma 

11, Torreya 26:91-93. 1926. isotyi 

'E. U.S.A. Florida. Bay Co.: 

ilunes along St. Ar 

idrews Ba 


>ster&RA. Matthews 12821; 

innotated as isotyj 

pe by D.E 

i. Dunn, 1969. 

Lythrum curtij 

isii Fernal 

id, Bot. Gaz. 33:155. 1902. isoTYPi 

E. U.S.A. Georgia. Calhoun 

:o.: miry place in 

swamp n 

earLeary, 20Augl901,A.H. Ov 

mss 6876; cited in original 

Marshallia obovata (Walt.) Beadle & RE. Boynton var. scaposa Channell, Contr. Gray 
lerb. 1 81 :90. 1957. isotype. U.S.A. North Carolina. Brunswick Co.: coarse sand, plowed 
re lane, 3 mi W of Leland along US 76, 2 1 May 1 949, R.K. Godfrey 49184; cited in original 
ublication, annotated as isotype by R.B. Channell, 1955 (holotype: GH). 

Mirabilis macfarlanei Const 



gon. Wall 

owa Co.: low< 

;r Cottonwood Landii 


en mouth of 

Somers Creek ; 


ittsburgh ; 

Landing, T27] 

M, R1-2W, 15 May 1^ 

?36, L. Cc 

mstance, R.C. 

Rollins, H.F. Cl 



ed in original 


Monarda X 


lioides W. 

'otmcan, Rhc 

)dora 61:303-304. IS 

)59. HOLO- 


Georgia. Un.o 


: highway 


SofBlairsville, 25Ju 

n 1959, \ 

V.H. Duncan 



TToc. Sci. Na 

Lt. Cherbourg 32:269. 

1901. ISO 


Florida. Swam 


aces in pin 

e barrens near 

Jacksonville, 10 Mar 

& 22 Oct 

1894, A.H. 


ted as isot) 


Jrd, 1968. 

Nuphar X 

interfluitans Fe 

.rnald, Rhodo 

ra 44:397-398. 1942 


U.S.A. ViR- 


It Co. 


, tidal water of 


idsor Shades, 

isotype by E.O. Beal, 1954 and J.H. Wiersma, 1' 
Oenothera HnifoliaNutt., J. Acad. Nat. Sci. P 

Georgia. DeKalb Co.: soil thinly overlying rock; 

A.H. Curttss 6778; annotated as isotype by G.B. ; 
Opuntia charlestonensis Clokey, Madrono 7:" 

Co.: Charleston Mountains, rocky slope, Gnffith 

Mill, rocky ndge, 13 Jul 1939, LW. Clokey 8430; cted in original publication, 

Phacelia dubia (L.) Trel. var. georgiana McVaugh, Ecol. Monogr. 13:158. 1943. 

ISOTYPE. U.S.A. Georgia. Oglethorpe Co.: shallow soil about granite outcrops, Echols Mill, 

12 mi NE of Lexington, 4 Apr 19^S, J.H. Pyron & R. McVaugh 2448; annotated as isotype 

by R. McVaugh, 1941. 

Physostegia serotina Shinners, Field & Lab. 24:17. 1956. isotype. U.S.A. Louisiana. 

Calcasieu Par.: clay ditch bank, E side of Lake Charles, 9 Oct 1955, L.H. Shinners 22108; 

Portulaca cornata Small, BuU.'Torrey Hot. Club 23:126-127. 1896. isotype. U.S.A. 
Georgia. DeKalb Co.: on Little Stone Mountain, 11 Sep 1894, J.K, Small s.n.; cited in 
original publication (holotype: NY). 

Portulaca smallii P. Wilson in Rydb., N Amer. El, 21:335. 1932. isotype. U.S.A. 
Georgia. DeKalb Co.: on Little Stone Mountain, 11 Sep 1894,/^. Small s.n.; cited in 
original publication (holotype: NY). 

PotentiUa beanii Clokey, Bull. S. Calif Acad. Sci. 38:4. 1939. isotypes. U.S.A. Nevada. 
Clark Co.: Charleston Mountains, ridge to Charleston Peak, 22 Jul 1938, /, W. Clokey 7974; 


res & R. Schult: 


eafl. : 

14:87. 1950. iSOTYPi 


Upper Rio Negro, Ig^irf 


ite mouth of R: 



.1942, R.E. 


nal publicatior 


.otype: GH). 

Primula nevadensi 


1967. isotyph. U.S. 

A. Nevada. 

White Pine Co.: Snake 

of Mount Was 

Georgia. DeKalb Co.: base ofLittle Stone Mountain, along Tom George Creek, 7 Jul 1893, 

Pycnanthemum curvipes (Greene) E. Grant & Epling, Leafl. Bot. Observ. 2: 140. 191 1 . 
isotype. U.S.A. Georgia. DeKalb Co.: upper slopes and summit of Stone Mountain, 27 Jul 
1895, J.K. Small s.n.; eked as type by E. Grant & C. Epling, 1943, Univ. Calif. Publ. Bot. 
20:209, annotated as isotype by J. Hamer, 1990 (holotype: US). 

Pyrrhopappus georgianus Shinners, Field & Lab 21:93. 1953. isotype. U.S.A. Geor- 

1947, A. Cronquist 4276; annotated as isotype by L.H. Shinners, 1953. 

Pyrus arbutifolia (L.) L.f. var. glabra Cronquist, Castanea 14:101-102. 1949. holo- 
type. U.S.A. Georgia. Wayne Co.: fine sandy soil in wet open pinelands SW of Jesup, 10 
Apr 1942, W.H. Duncan 4659; cited in original publication. 

Co.: wash in main fork of Alamo Canyon, Ajo Mountains, Organ Pipe Cactus 
Monument, 15 Aug 1952, C.H. Muller 9^19; cited in original publication. 

Quercus oglethorpensis W. Duncan, Amer. Midi. Naturalist 24:755. 1940 
U.S.A. Georgia. Oglethorpe Co.: well drained bottomland of Buffalo Creek 6 
Lexington, 1 1 Sep 1940, W.H. Duncan 2870; cited in original publication (holot 

Sanicula deserticola C.R. Bell, Univ. Calif. Publ. Bot. 27:133-230. 1954 
MEXICO. Baja California: Norte, rocky slopes 6 mi N of American Smelter Co 
Feb 1953, C.R. Bell 1355; cited in original publication. 

Senecio andersonii Clokey, Bull. S. Calif. Acad. Sci. 37:10. 1938. isotyp: 
Nevada. Clark Co.: Charleston Mountains, Lee Canyon, brushy meadow, yellow | 
5 Aug 1935, l.W. Clokey 5638; cited in original publication. 

Senecio ganderi T.M. Barkley & R.M. Beauch., Brittonia 26:106-108. 1974 
U.S.A. California: San Diego Co.: N slope of SE ridge, Lawson Peak, 23 May 
Moran & M. Douglas 21038; cited in original publication (holotype: NY). 

Silene andersonii Clokey, Bull. S. Calif. Acad. Sci. 38:2. 1939. isotype. U.S.A. 
Clark Co.: Charleston Mountains, Levell Canyon, steep N gravelly slope, 9 Aug 1 

Solidago rigida L. var. laevicaulis Shinners, Field &Lab. 19:35. 1951. isotyp 
Texas. Dallas Co.: 2 mi N of Cedar Hill, limestone soil, 11 Nov 1947, £. Whitehou 
cited in original publication, annotated as isotype by S. Heard, 1986 (holotype 

Solidago tarda Mackenzie ex Small, Man. SE Fl. 1355, 1509. 1933. isotyp 
Georgia. Clarke Co.: Winterville, Hulme Farm, 20 Oct 1926, J. H. Miller s.n. 

danis holzinger McVaugh, Wrightia 1:45-46. 1945. isotype. U.S.A. Oklahoma. 
n Co.: open prairie near Washington, 12 Jun 1936, M. Hopkins &C.T. Eskew 688; 

ition, annotated as isotype by K.L. Wilbur, lyby 
■ade, Torreya 31:71. 1931. isotype. U.S.A. Florie 

type by W.H. Can 

ip, 1942. 


ii Chapm., Fl. S U.S. 260. I860, isotype? U.S.A. Florida: 


zn s.n.; annotated as isotype? By S. McDaniel, 1986. 

:umBuckley,Amer.J.Sci.45:175. 1843. isotype? U.S.^ 

s. ssp. longistylis S.C. Keeley, J. Arnold 
: Basse Terre, limestone cliffs along road; 
along highway to St. Ann, 20 Dec 197 ^ J. E. Keeley & S 

Vernonia angusi 

tifolia Michaux var. i 

rnohrii S.B. Jones, Rhodoraf 


holotype. U.S.A. A 

[A. Washington 

Co.: roads 





m I406;c. 

ited in original pub 

4ication, anr 

as holotype by S.B.. 


. 1964. 

; McVaugh, Cc 

Michigan Herb. 

isotype. MEXICO.] 


): mountains 3 mi S of La Huerta, road to Barra de Navidad 


; annotated as i; 

iotype by 5 

5.B. Jones, 1972. 

Vernonia duncanii S 

!b. Jones, Brit 

toma 25:1 

08-109. 1973. Hc 

)lotype & IS 


nestone hillside 


W of Cuidad Victor 

■ia, km 155,1 

Hwy 101, 6 Nov 1970, S.B. Junes 20539, cited in original publication, annotated as 

Vernonia harleyi H. Robinson, Phytologia 44 287-288 1979 isotype. BRAZIL. 
Bahia: 19.5 km SE of town of Morro do Chapeu on road BA052 to Mundo Novo, by the 
Rio Ferro Doido, 2 Mar 1977, R.M. Hariey 19296; cited in original publication, annotated 
as isotype by J.G. Stutts, 1980 (holotype: US). 

Vernonialiatrioicies DC ssp.gentryiS.B.Jones,Rhodora 78: 193-194. 1976. HOLOTYPE 
& ISOTYPE. MEXICO. DURANGO: 1 16 mi W of Durango on Hv^y 40, 7 Mar 1974, S.B.Jones 
22527; cited in original publication, annotated as holotype and isotype by S.B.Jones, 1974. 

Vernonialibertadensis S.B. Jones, Fieldiana,Bot. 5:3 1.1980. HOLOTYPE. PERU: Depto 
La Libertad: Prov. Otuzco, Cerro Sango, 7 Jun 1953, A. Lopez 1947; cited in original 
publication, annotated as holotype by S.B. Jones, 1978. 

Vernonia macphersonii S.B. Jones & J.G. Stutts, Brittonia 33:546. 1981. isotype. 
MEXICO. Jalisco: steep mountainsides 3-10 km generally E on the road to Mina del Cuale, 
from the jet. 5 km NWof El Tuito, Mpio. de Cabo Corrientes, 16-19 Feb 1975, ^. MfV^/^^/- 
26442; cited in original publication (holotype: MICH). 

Vernonia macvaughii S.B. Jones, Brirtonia 25:105. 1973. holotype. MEXICO. 
Oaxaca: 4.1 mi S of Puebla-Oaxaca State line and 6 mi N of Huajuapan de Leon, 23 Mar 


Vernonia michoacana McVaugh, Contr. Univ. Michigan Herb. 9:482-484. 1972. 
isotype. MEXICO. Michoacan: steep dry mountainsides ca. 8 km NW of Aguililla, road to 
Aserradero Dos Aguas, 2 Mar 1965, i?, AlrV^a^>?>2266S;annotatedasisotypeby S.B. Jones, 

Vernonia nestor S. Moore, J. Linn. Soc, Bot. 35:317. 1902. isotype. MALAWI: 
Nyasaland, 1895,/ Buchanan 129; cited in original publication. 

Vernonia obtusa (Gleason) S.R Blake ssp. parkeri S.B.Jones, Brittonia 25:108. 1973. 
holotype, MEXICO. Hidalgo: dry limestone hillside 4-5 mi SW of Jalaca at La Placita, 7 
Nov 1970, S.B.Jones 20564; cited in original publication, annotated as holotype by S.B. 
Jones, 1972. 

Vernonia oligactoides Less., Linnaea VI;648. 1831. neotype. BRAZIL: Sao Paulo, 
Mun. Itarare, PRl 51, Fazenda Esplanada, 17 Feb 1982,7.G, Siutts 1214, cited as noeotype 
by J.G. Stutts, 1988, Rhodora 90:86. 

Vernonia paucartambensis Dillon, Brittonia 36:336. 1984. isotype. PERU: Cuzco, 
Paucartambo, km 132 on Paucartambo-Pilcopata Road, 5 km below PiUahuata, 27 Jun 
1978, A. Gentry, M. Dillon, P. Berry & J. Aronson 23553; cited in original publication 

Vernonia pugana S.B. Jones & J.G. Stutts, Brittonia 33:544-545. 1981. 
MEXICO. Jalisco: foothills of Sierra de Manantlan, 16-22 km S of El Chante, ; 
1975, R. McVaugh 26172; cited in original publication (holotype: MICH). 

Vernonia salicifolia (DC) Schultz-Bip. var. baadii McVaugh, Contr. Univ. M 
Herb. 9:484. isotype. MEXICO. Jalisco: headwaters of Rio Mascota, 8-10 km 
Rincon, on the road to Aserradero La Cumbre, 2-3 Apr 1 965 , i?. McVaugh 23473;-s.n 
as isotype by S.B. Jones, 1972. 

Moore and Giannasi, Type specimens, University of Georgia herbarium 151 


Vernonia woytkowskii S.B. Jones, Fieldiana, Bot. 5:29. 1980. isotype. PERU: 
Lambayeque, Purculla ad Olmos, 29 Sep 1961, F. Woytkowski 6770; cited in original 
publication, annotated as isotype by S.B. Jones, 1978 (holotype: MO). 

Viola charlestonensis Baker & R.T. Clausen, Madroiio 8:58. 1945. isotype. U.S.A. 
Nevada. Clark Co.: Charleston Mountains, Forest Camp No. 1, ridge S of Deer Creek, 23 
May 1937, l.W. Clokey 7301; cited in origmal publication. 

Vitis acerifolia Raf., Amer. man. grape vines 14. 1830. neotype. U.S.A. Texas. 
Wilbarger Co.: along Beaver Creek on US 283 S of Vernon, in rest area 1.5 mi S of jet. with 
Farm Road 1763, 13Jun 1986, M.O. Moori? 700; cited as neotype by M.O. Moore, 1991, Sida 

Vitis argentifolia T.V Munson, Proc. Soc. Promot. Agric. Sci. 8:59- 1887. neotype. 
U.S.A. West Virginia. Nicholas Co.: W side of US 1 9, 1 .7 mi S of the Braxton Co. line, 12.3 
mi N of jet. with WV 55, 25 Aug 1987, M.O. Moore 886; cited as neotype by M.O. Moore, 
1991, Sida 14:347-348. 

Vitis bloodworthiana Comeaux, Sida 14:460-461. 1991. isotype. MEXICO. Du- 
RANGO: along Hwy 40 1 6.5 km W of Del Diablo and 35.5 km E of the tropic of cancer, 1 
Jul 1985, B.L. Comeaux 4219; cited in original publication (holotype: SMU). 

Zornia bracteata (Walter) J.F. Gmel., Syst. Nat.:1096. 1791- isoneotype. U.S.A. 
Georgia. McDuffie Co.: open pine hickory woods 5 mi E, 43° south of Thomson, 29 Jul 
1950, W.H. Duncan 11337; cited as isoneotype by R.H. Mohienbrock, I96I, Webbia 
16:29-30 (neotype: US). 

N.C. and S.B. Jones. 1985. Georgia plant list. Second edition. Departn- 
S.B. and N.C. Coile. 1979a. Plant list for Clarke County, Georgia. I 

S.B. and N.C. Coile. 1979b. Georgia plant list. Department of Botai 
Georgia. Athens. 

S.B. and N.C. Coile. 1988. The distribution of the vascular flor 
jartment of Botany, University of Georgia. Athens. 


Taylor, Walter Kinglsey. 1992. The Guide to Florida Wildflowers. 
(ISBN 0-87833-747-4, hbk) Taylor Publishing Company, 1550 West 
Mockingbird Lane, Dallas, TX 75235. $24.95. 320 pp. 
Are you headed to Florida this year and would like to identify some of the roadside flowers 

book is designed be a pictorial guide to many of the commonly occurring wildflowers, 
especially of central and northern Florida. Taylor's prowess as a photographer and enthusi- 
astic field botanist comes through clearly, as the user will find a usually high quality color 
photograph of each species discussed. The photos are further supported by thumbnail 
descriptions, flowering times and geographic occurrences within the state. Conspicuous 
fruits are shown as photo insets for a number of the species, as well. However, close-ups of 
some with small, diffusely arranged flowers would have been a welcomed addition. 

One's first impression, based on the crisp layout and handsome photos, is that this book 
is a user-friendly guide; frustration lies ahead. Species are arranged first by color (color tabs 
at the top of each page), and secondly by an outdated sequence of relationships. Thus, one 
is guided to the species by looking through all the photographs of a particular color group. 

his task is CO 


finition of col 

or groups. That is 

, should : 

irticular specii 

irched for in the pinko 

,r the violet gn 



een group 

yellow ; 

oice IS not alw 

ays obvious because 

• the v.olei 

roup contains 

ik ones. Furthermore, . 

n the yellc 

,w group 1 



light yellow petals are i 



out picture 

:s showing 

hite petals. 

Florida alwa) 

^s prese 

nts a problem to desig 

;ners of plant. 


tion manu 

als. In th( 


ided frc 

s, there IS a myriad of d 
>m more tropical areas, 

iverse tropical 



only therf 
. Nationa 

aides usually i 

gnore all of the tropical species, while state i 

=^uides St, 


1 which tc 


; reason 

.visitors traveling to South Florida w 


trated furt 



ing om 


a, a prominent 


A flower ol 

F roadside; 

1 not only Soui 

)f central Flori 

11. The aut 

; excused base. 

d on his 

; statement, "...exclude 

d for the most 

part the 

woody plants, excep, 

T some trees, s 


and vines that are rath 

er common an 

d produc 

e flowers.' 

i furtherm( 


iclude the mo 

re nort 

hern and less showy ; 

.crub plum, n 

sd choke 

berry, gopher apple 

-eenbners, Ite^ 

are conspicuoi 




aesalpima bona 

luc, Car 

■tea, Echites, Epidendrm 


ncidium, Ricmu 

is, Thai 

ia, Tillandsia, Urechites 

, and Waltheric 


Although thi; 


.as several problems, it i 

s certainly no worse thai 

1 other Sim 

ilar guide: is 

well wc 

irth the money, if you do not already o 

wn one of 

•the others 

guides anc 

)u do not expect to 1 

3e guided through tropical Florida.- 

-Roger ' 

W. Sanders, 


isearch Institute 

'Of Texas. 





North Carolina Natural Heritage Program 

132 N orris Road 

Swamboro, NC 28584, U.S.A. 


North Carolina Natural Heritage Program 

Department of Environment, Health, and Natural Resources 

P.O. Box 27687 

Raleigh, NC 27611-7687 , U.S. A 

University of Michigan Herbarium 

North University Building 

Ann Arbor, Ml 48109, U.S.A. 


Ontario Ministry of Natural Resources 

P.O. Box 9000 

Huntsville, Ontario, POA IKO, CANADA 

Carex lutea (section Ceratocystts) 

is described from th 

e outer coastal plain of so 

North Carolina. It is the only specie 

cur south of New Jersey ak 

Atlantic Coast, and appears to be a lo^ 

calized endemic of we 

;t savannas underlam by Im 

deposits. From all other species oft 

his section, C. lutea i 

s distinguished by its pale 

:ulms, and elongate i 


long with usually only one or two 

widely separated pi 

stiUate spikes below the 

Field surveys in 1991 by R.J. LeBlond of a i 
nderlain by coquina limestone deposits on the ou 

154 S]DA16(1) 1994 

County, North Carolina disclosed an unfamiliar Carex with clear affinity to 
the well-defined boreal section Ceratocystis, and more specifically with the C. 
flava L. complex as defined by Crins & Ball (1988, 1989a, 1989b). Sub- 
sequent searches for the plant by LeBlond and A.S. Weakley disclosed four 
additional populations in Onslow and Pender counties. However, the popu- 
lations were all clustered within a radius of about 4 km. Furthermore, only 
the original population was large; all the others consisted of fewer than 60 
clumps each. 

Diagnostic characters of the Carex flava complex include globose to ovoid, 
more or less approximate, short-peduncled to sessile pistillate spikes with 
the perigynia spreading (the lowermost often refiexed) (Crins & Ball 1 989b). 
Three species of the complex, C. cryptolepis Mack., C. flava, and C. vtridula 
Michx. subsp. viridula, are known from the northern coastal plain, but none 
has been found south of northern New Jersey (Crins & Ball 1989b). Two 
additional subspecies of C. vtridula, C. viridula subsp. oedocarpa (Andersson) 
B. Schmid and subsp. brachyrrhyncha (Celak.) B. Schmid, are known from 
boreal habitats farther north along the Atlantic coast in the Gulf of St. 
Lawrence region and in Eurasia. Several other, distinct species are found 
elsewhere in the old world and in the temperate parts of the southern 
hemisphere, but the relationships of the North Carolina plants are clearly 
with the North American members of the C. flava complex. 

Close examination of the North Carolina plant discloses a number of 
significant points of difference from the other species of the Carex flava 
complex in North America. The pistillate scales of the North Carolina plant 
are pale yellowish-green, immediately separating it from C. flava and most 
subspecies of C, viridula, the scales of which are coppery or tinged with red 
or brown. The larger (up to 5.2 (5.8) mm long), reflexed lower perigynia 
separate it from those subspecies of C, viridula that may have paler scales. 
The North Carolina plants can be separated from the pale-scaled C. cryptolepis 
by its usually sparsely serrulate perigynium beaks, as well as its taller stature 
and longer inflorescences. 

Several features unique to the North Carolina plant separate it from all 
other species in the C. flava complex both in North America and world-wide. 
The North Carolina plant is taller than any other members of the section, with 
taller mature flowering individuals typically over 65 cm tall and reaching 
heights of up to 1.25 m. Even though the plants are tall, the number of pis- 
tillate spikes is normally only one or two, extremely rarely three. As well, the 
inflorescences of the North Carolina plants (particularly those of culms that 
produce two pistillate spikes) are much longer than any other species, the 
longest ranging from 18-41 cm. All other members of the C. flava complex 
are much shorter, rarely reaching heights of over 65 cm, and routinely pro- 
ducing 2-3 pistillate spikes in shorter inflorescences 1.5-12(20) cm long. 

Based on its unique morphology, this rare and highly localized North 
Carolina plant is here described as a new species. 

Carex lutea LeBlond, sp. nov. (section Ceratocystis). (Fig. 1) 

Styli marcesentes; stigmata 3. Antherae 3, 2.1-3.6 mm longae. 

Plants cespitose in small to large (up to ca. 45 fertile culms) clumps, with 
short ascending rhizomes; roots pale brown, not densely felted with root 
hairs; fertile culms (40)65-110(125) cm tall, 0.6-0.9 mm wide at base of 
lowermost spike, central, more or less trigonous, smooth except at apex 
where often slightly scabrous-angled, with glabrous, stramineous to pale 
brown bladeless basal sheaths. Leaves 3-7, mostly basal; blades 5.5-28 cm 
long, much shorter than culms, 1 .8-3.8 mm wide, plicate, yellowish-green, 
glabrous, the margins antrorsely scabrous, the widest leaves 2.4-3.8 mm 
wide; leaf sheaths 2.5-11 mm long, more or less tightly enveloping culms, 
glabrous, green; inner band of sheath glabrous, whitish-hyaline, the apex 
thin and friable, irregularly concave, more or less truncate, or occasionally 
slightly prolonged; ligules obtuse, 1.4-3.6 mm long, the free portion more 
or less entire, whitish-hyaline, to 0.7 mm long. Vegetative shoots 40-65(90) 
cm tall; leaves 5-11, similar to those of fertile culms but up to 65(90) cm 
long and 4.6 mm wide, more or less evergreen (at least the proximal portions 
of the blades); pseudoculms ca. 5-13 cm tall. Inflorescences (4.5)5.4-25(41) 
cm long, with all spikes quite separate, the lowest 2 pistillate spikes (if 
present) (1.6)4.5-18(33) cm distant; spikes single at nodes, ascending; 
lowermost spikes with peduncles 0.4-4.5(16.5) cm long, the uppermost 
pistillate spike sessile; peduncles smooth and terete proximally, more or less 
trigonous and serrulate-angled distally; lowermost bracts usually reflexed 
(except when subtending long-peduncled spikes), with blades 5-2 5 cm long 
and 1.2-2.5 mm wide and sheaths (0)0.2-4 cm long, 0.5-1.3(1.9) times as 
long as the inflorescence, the uppermost bracts also reflexed, bur much 
reduced. Spikes 2-3(4), the terminal staminate (very rarely with an addi- 
tional small, accessory staminate spike at base), the lateral pistillate (very 
rarely with a small staminate apex up to 8 mm long). Terminal spikes (9) 1 7- 
39 mm long, 1.4-2.5 mm wide, ca. 45-90-flowered, peduncles (0.3)1- 
6(10. 2)cm long, (0.1)0.7-2. 5(5.1) times as longas the spikes. Lateral spikes 
0.7-2.7 cm long, 8-11 mm wide, globose to ellipsoid or short-cylindric. 

LeBlond, Weakley, Reznicek, CRINS, Carex lutea 157 

densely 15-60-flowered. Pistillate scales 2.1-3.3 mm long, 0.9-1.3 mm 
wide, lanceolate to narrowly ovate, obtuse to more or less acuminate, 
glabrous, pale yellowish-green with a green center and wide hyaline 
(sometimes stramineous-tinged) margins and apex, 1 -nerved, almost totally 
concealed by the crowded perigynia. Staminate scales 2.9-5.6 mm long, 
1.2-1.9 mm wide, narrowly obovate to narrowly oblong, obtuse to acute, 
glabrous, stramineous to pale brown with a green center and wide hyaline 
margins and apex, l(3)-nerved. Perigynia (3.5)3.9-5.2(5.8) mm long, 1.4- 
2.2 mm wide, spreading, strongly outcurved and squarrose in the spikes, the 
lowermost strongly reflexed, somewhat inflated, irregularly and asymmetri- 
cally trigonous with convex, obovate sides, glabrous, bright yellow proxi- 
mally, pale yellowish-green distally, short-stipitate and tapered to the base, 
7-13-nerved with two nerves much more prominent than the rest, con- 
tracted into a strongly deflexed beak; beaks 1.4-2.2 mm long, finely and 
sparsely serrulate or sometimes smooth, green, the apex bidentulate with 
teeth 0.2-0.5 mm long. Achenes 1.4-1.9 mm long, 1.2-1.5 mm wide, 
trigonous with flat to slightly concave, strongly obovate sides, essentially 
truncate apically, brown, sessile. Styles withering; stigmas 3. Anthers 3, 

Type: UNITED STATES. North Carolina. Pender Co.: SW of N.C. Hwy 50, ca. 1,8 mi 
WofOnslowCo. line, ca. 11 mi NNW of Holly Ridge. S of secondary Rd 1532,0.85 mi 
SW of N.C. Hwy 50, "Lamer Quarry Savanna," 20 May 1993, A.A. Reznicek 8942 with S.A. 
Rezmcek. R.J. LeBlond & B.A. Some (holotype: MICH; isotypes: BRIT/SMU, FLAS, GA, 
NCU, TAES, us, VDB, herb. C.T. Bryson). 

Addirional specimens examined: NORTH CAROLINA. Onslow Co.: NE side N.C. 
Hwy 50, ca. 10.4 mi NNW of Holly Ridge, "Powerline Savanna," 20 May 1992, A.A. 
Reznicek 8954 with S.A. Rezmcek, R.J. LeBlond, & B.A. Sorrie (MICH, VPI). Pender Co.: 
Lanier Quarry Savanna, S of secondary road 1532 0.85 mi SW of Hwy NC 50, 1 1 Apr 1990, 
R.J. LeBlond 1188 (NCU); 24 Apr 1991, RJ- LeBlond 1983 (NCU); 22 May 1991, R-J- 
LeBlond 2056 (MICH, NCU); 27 Jun 1991, RJ. LeBlond 2267 (NCU); 20 Apr 1992, R.J. 

LeBlond 2723 

(NCU); Sandy Run Savanna 

along Hunt Club Rd ( 

).5 miS 


12 May 1992. 

, A.S. Weakley & R.J. LeBlond s 


Lvanna, 1 


N.C. 50ca. 

ne, 12 May 1992, A.^ 

. Weakle 

y & R.J LeBlond 





a NNW of Holly 

Ridge, "Wacl 

cms Savanna," 20 May 1992 

, A.A. Rezmcek 8949 

with S.. 

A. Reznicek, R.J. 

LeBlond &B.. 

\. Sorrie (MICH, USCH). 


In the key to the Carex flava complex in Crins & Ball (1 989b), C. lutea keys 
closest to C. cryptolepis because of its pale, yellowish-green scales, narrow 
leaves, and perigynia of similar size. The overall yellowish-green color of C. 
lutea is also similar to that of C cryptolepis. Insertion of the following couplet 
in place of the first lead of couplet 3, however, will complete the key, and 

allow easy separation ot C. lutea Irom C. cryptoLepis. 

times as long as the inflorescence; staminate spike peduncles mostly 0.7-2.5 
times as long as the staminate spikes; achenes 1.2-1.5 mm wide; perigynium 
beaks often sparsely serrulate C. lutea 

as long as the inflorescence; staminate spike peduncles 0.2-0.5 times as long 
C. cryptolepis 

Although Carex lutea keys with C. cryptolepis due to its pale scales, its 
elongate inflorescence is most similar in aspect to that of C. vindula subsp. 
brachyrrhyncha var. elatior{^ch.\td\.) Crins (C. lepidocarpa Tausch), a boreal and 
subarctic taxon of highly calcareous, open wetlands in the Gulf of St. 
Lawrence region and Europe. Carex lutea differs most obviously from C. 
viridula subsp. brachyrrhyncha var. elatior in plant size, inflorescence size, and 
paler scales. The two taxa can be separated by the following couplet. 
1. Pistillate scales pale yellowish-green, nearly the same color as the perigynia 

and thus inconspicuous; tallest culms 65-1 10(125) cm; longest inflorescences 

1. Pistillate scales with brown coloration, conspicuous among the greenish to 
yellowish perigynia; tallest culms (10)20-60(85) cm; longest inflorescences 

2-12(18) cm long C. vindula subsp. brachyrrhyncha var. elatior 

In addition to the scale color and size differences, Carex lutea differs from 
C. viridula subsp. brachyrrhyncha var. elatior in inflorescence proportions. In 
C. lutea, the bracts of the lowest pistillate spikes are shorter than or equalling 
the inflorescence and most staminate spikes are shorter than their peduncles. 
In C. viridula subsp. brachyrrhyncha var. elatior, the bracts of the lowest 
pistillate spikes often equal or exceed the inflorescence and most staminate 
spikes are longer than their peduncles. Because of the great variability in 
these proportions due to differences in the number of pistillate spikes in the 
inflorescence and the considerable plasticity of inflorescences depending on 
the vigor of the plants, there is modest overlap in these proportions, 
rendering them unsuitable as key characters. Carex lutea also has proportion- 
ately narrower leaves than C. viridula subsp. brachyrrhyncha var. elatior. 
Although a significantly shorter plant than C. lutea, the widest leaves of C. 
viridula subsp. brachyrrhyncha var. elatior range up to 5.6 mm whereas those 
of C, lutea are at most 4.6 mm wide. Other differences include the larger 
staminate spikes of C lutea, which are up to 39 mm long, much larger than 
the maximum of 25 mm in C. viridula subsp. brachyrrhyncha var. elatior, and 
larger perigynia up to 5.2(5.8) mm long in C. lutea whereas those of C. 
viridula subsp. brachyrrhyncha var. elatior are only up to 4.2 mm long. 

It is tempting to suggest that the character of pale scales, which uniquely 
defines the two North American endemics, C. cryptolepis and C. lutea, indi- 

LeBlond, Weakley, Reznicek, Crins, Carex lutea 159 

cates a close relationship. However, until genetic, breeding system, and 
micromorphological evidence comparable to that available for the other taxa 
is known for C. lutea, statements of putative relationship are speculative. 
Nevertheless, when a character compatibility analysis was run with C. lutea 
added to the data matrix of Crms ( 1 990), C. lutea consistently clustered with 
C. cryptolepis (unpublished data). Although genetic and micromorphological 
data are not yet available for C. lutea, it can be recognized readily and 

The ecology of C. lutea is quite distinctive within the section; not 
surprisingly, since its occurrence is so far removed from all the other species. 
All the sites occur in sandy soils overlying coquina limestone deposits, and 
the species shows a preference for the ecotone between the longleaf pine 
savanna and nonriverine swamp forest communities. Soil from a Lanier 
Quarry microsite for C. lutea had a pH of 5.6. This is similar to pH levels 
recorded at Thalictrum cooleyi H.E. Ahles microsites at the Lanier Quarry 
Savanna. Thalictrum cooleyi is the most frequent associate of C. lutea, and is 
similarly restricted in distribution. Soils not supporting T. cooleyi at the 
Lanier Quarry site regularly test at lower pH levels. 

Most Carex lutea plants occur in the partially to moderately densely tree 
shaded savanna-swamp ecotone, with scattered shrubs and a moderate to 
dense herb layer. The savanna-swamp ecotone is subject to occasional to 
frequent fires which favor a herbaceous ground layer and suppress shrub 
dominance. Carex lutea is a subdominant to patch dominant at two of the five 
known sites. Occurrences are densest in areas of partial to moderate tree 
shading with an absence of a shrub understory. Frequent associates include 
Taxodium ascendens Brongn., Liriodendron tulipifera L., Acer rubrum L., Myrica 
cerifera L. var. cerifera, Thalictrum cooleyi, Aletris farinosa L., Carex lonchocarpa 
Willd., Osmunda regalis L. var. spectabilis (Willd.) A. Gray, Physostegia purpurea 
(Walter) S.F. Blake, and Parnassia caroliniana Michx. 

Though the height oi Carex lutea suggests that it might be easily found, 
the plants are slender, occur in areas with substantial graminoid cover, and, 
except for one site, are rare and localized. Thus, a colony may remain quite 
inconspicuous, even in fruit. This suggests the possibility that additional 

coastal plain, though the association with narrowly endemic species in an 
unusual habitat suggests that C. lutea may be a highly localized species. 

The occurrence of a localized disjunct species of the boreal Carex flava 
complex on the outer coastal plain of North Carolina is quite striking. This 
constitutes a disjunction of about 750 km from the nearest known popula- 
tions of other members of sect. Ceratocystis. In Eurasia, species of this com- 
plex occur as far south as Spain, Morocco, Turkey, and Iran. The most sub- 
stantial range separation among close relatives in Europe is that between C. 

vlridula subsp. brachyrrhyncha vars. elatior and nevadensis (Boiss. & Reut.) 
Crins from southeastern France to southern Spain (ca. 800 km). 

The type locaHty of Carex lutea, Lanier Quarry Savanna, and the other 
locations where it has been found are ecologically highly unusual and phyto- 
geographically notable. The combination of fairly open conditions under- 
lain by a calcareous substrate is very rare on the Atlantic coastal plain. Over 
26 species regarded as rare in North Carolina by the state Natural Heritage 
Program are found at the Lanier Quarry site. Nine of these are listed as 
endangered, threatened or candidate species by the U.S. Fish and Wildlife 
service. Many of these rare plants have very restricted distributions, either 
being endemic to small areas or with highly scattered occurrences. The 
affinities of these taxa are variable, but include connections to the calcareous 
savannas of the Gulf Coast states, alkaline marshes of the Atlantic tidewater, 
calcareous glades, barrens, and prairies of the Appalachian region, and 
pinelands of the Carolinas and southern New Jersey. Most notable are the 
two endemic species closely associated with Carex lutea, Thalictrum cooleyi 
and an undescribed species oi Allium. Thalictrum cooleyi is endemic to about 
a dozen sites, all savanna margins in sites underlain by calcareous deposits, 
in North Carolina, Georgia, and Florida. It has the highest ploidy level 
known in the genus (30x at2n = 2 10), suggesting its derivation from a more 
widespread northern and inland species such as T. revolutum (Park 1 992). The 
undescribed Allium is related to the widespread inland species A. cernuum 
Roth and A. stellatum Ker and is known from only 5 sites, four of them shared 
with C. lutea. 

calcareous deposits have evidently served as a small archipelago of phytogeo- 
graphic islands for species poorly adapted to present conditions on the 
southeastern coastal plain. It seems likely that Carex lutea is a narrowly 
distributed, very rare endemic. It may reflect a southern extension of the C. 
flava complex during the Pleistocene glaciation, followed by isolation in a 
few suitable sites and speciation. Alternatively, C. lutea could represent the 
relictual occurrence of a formerly more widespread and older species in the 
C/^m complex. 

We are grateful to the North Carolma chapter of The Natu 
for providing funding for a systematic survey of the flora of it 
preserve, which resulted in the discovery of this new specie 

LeBlond, Weakley, Reznicek, Crins, Carex lutea 

Crins, WJ. and P. W. Ball. 1 988. Sectional limits and phylogenetic considerations 
section Ceratocystis (Cyperaceae). Brittonia 40:38-47. 

North America and northern Eurasia. I. Numerical taxonomy and character analy 
J. Bot. 67:1032-1047. 
Crins, W.J. and P.W. Ball. 1989b. Taxonomy of the Carex fiava complex (Cyperj 

Park, M.M. 1992. A biosystematic study oiThalictrum section Leucocoma (Ranunci 


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S.M. Tracy Herbarium 

Department of Rangeland Ecology and Management 

Texas A&M University 

College Station, TX 77845-2126, U.S.A. 

Sporobolus potosiensis is a new rhizomatous species from San Luis Potosi, Mexico that dil 
rom its closest putative relative, S. airoides, by 1 .) having conspicuous and slender rhizor 
:.) lacking trichomes behind the ligule, and 3.) generally smaller in stature. A 
ombination is proposed at the subspecific rank: S. airoides subsp. regis. A key to 
hizomatous species of Mexican Sporobolus, as well as S. nealleyi and 5". airoides, is provic 

Sporolobus potosiensis es una nueva especie rizomatosa de San Luis Potosi, Mexico, que 
difiere de su pariente putative mas proximo S. airoides por 1 .) tener rizomas conspicuos y mas 
delgados, 2 .) ausencia de tricomas detras de la ligula, y 3 .) tamano mas pequeno generalmente. 
Se propone una nueva combinacion de rango subespecifico: S. airoides subsp. regis. Se ofrece 

Sporobolus is a genus of approximately 1 60 species, distributed throughout 
the tropics, subtropics and temperate areas (Clayton and Renoize 1 986). The 
species of Sporobolus intergrade to such an extent that their limits are seldom 
sharply defined (Clayton and Renvoize 1986). There are approximately 60 
species of Sporobolus known to occur in the New World (Judziewicz and 
Peterson 1989) and 28 species are reported from Mexico (Beetle 1987), only 
two species of which have been reported as rhizomatous: -S". virginicus (L.) 
Kunth and S. regis I.M. Johnston. Sporobolus virginicus, a strongly rhizoma- 
tous perennial, is found on sandy beaches and at the bases of sand dunes from 
Virginia to Texas, south to Brazil and Peru, the West Indies, and the tropics 
of the Old World (Reeder 1975; Pohl 1980). 

Sporobolus regis is only known from Coahuila, Mexico (Beetle 1987). 
Johnston (1943) stated that his new species, S. regis, was probably most 
closely related to 5'. airoides (Torr.) Torr. and 5". wrightii Munro ex Lamson- 
Scribner and differed from these species in having pubescent leaf sheaths, 
tufts of trichomes in the axils of the panicle branches, and very coarse 
SiDA 16(1): 163- 169. 1994 

164 Sid A 16(1) 1994 

rhizomes. Johnston (1943) also mentioned that "the bases of the culms and 
the younger nodes of the rhizomes bear shredded remnants of old leaves." 
The presence of shredded remnants of old leaves on the rhizomes needed 
further investigation, since grass rhizomes usually produce modified leaves 
termed "scale leaves" (Gould and Shaw 1983). Examination of the holotype 
{Stewart 2655 (GH, acronyms according to Holmgren et al. 1990)}, revealed 
that S. regis is not rhizomatous and that the "rhizomes" are the portion of the 
culms covered by soil, with their sheaths decomposed and their nodes 
developing adventitious roots, thus resembling rhizomes. All of the tillers, 
or shoots, of the type specimen are erect and lack lateral (horizontal) shoots. 
Because of the obvious remnants of the culm sheaths, and the lack of "scale 
leaves" on the culms, there is little doubt that the "rhizomes" of 5". regis are 
in fact culms responding to being buried. 

Sporobolus regis is distinguished from S. airoides by its densely pubescent 
sheaths and tuft of trichomes in the axils of the panicle branches. In our 
opinion, these characters are not significant to warrant the recognition of cS^. 
regis at the specific rank, but are significant to warrant infraspecific recog- 
nition. The lack of rhizomes places this taxon in S. airoides. However, these 
unique and distinct characters coupled with its restricted distribution (SW 
Coahuila) justify the recognition of this taxon at the subspecific rank, thus 
necessitating the following new combination. 

Sporobolus airoides (Torr.) Torr. subsp. regis (I.M. Johnst.) Wipff & S.D. 
Jones, comb, et stat. nov. basionym: Sporobolus regis I.M Johnst., J. Arnold Arbor. 
24: 393-394 (1943). Type: MEXICO. Coahuila: salt flat 4 km SE of Laguna del Rey, 

Sporobolus potosiensts^ipii Sc S.D.Jones is the second rhizomatous species 
known to occur in Mexico. This new species was originally identified at 
TABS from specimens that were either unidentified or misidentified as S. 
nealleyi Vasey. Specimens were requested from MEXU, MICH, MO, TEX 
and US to determine whether additional collections could be found. 

Sporobolus nealleyi is a cespitose, non-rhizomatous, gypsophilous species of 
the southwestern United States and northern Mexico. Chase (1951) and 
Correll and Johnston (1970) reported S. nealleyi as rhizomatous or subrhi- 
zomatous. None of the specimens examined, including the holotype {Nealley, 
US), had rhizomes. However, the densely tufted and persistent bases can be 
covered by soil and then appear to be shortly rhizomatous or subrhizoma- 
tous, as already discussed above. Sporobolus nealleyi, in Mexico, is reported 
from Coahuila, Nuevo Leon, and San Luis Potosi (Reeder 1975; Beetle 
1987). It appears that reports from San Luis Potosi are based on misidenti- 
fied collections of S. potosiensis. Based upon specimens examined, the 
distribution of S. nealleyi in Mexico is probably restricted to northern 
Coahuila and Nuevo Leon. 

The closest putative relative oi S . potosiensis is S. airoides. Specimens of 5". 
potostensis superficially resemble depauperate individuals of .S. airoides, but 
differ from S. airoides by 1.) having conspicuous and slender rhizomes, 2.) 
lacking trichomes behind the ligule, and 3.) being smaller in stature. 
Sporobolus airoides does not have rhizomes and usually has conspicuous 
trichomes behind the ligule, though some very depauperate specimens were 
examined that did not have trichomes. Sporobolus airoides is usually a robust 
plant to 150 cm tall, but depauperate specimens may resemble S . potosiensis 
in general appearance, except for the conspicuous rhizomes. 

2. Inflorescences (5-) 15-25 cm wide, foLuid on dry soils in open ground, 

2. Inflorescences 1-3 5 cm wide, restncied to gypsifeious soils S nealltyi Vasey 

(I.M. Johnst.) Wipff & S.D. Jones 
3 . Back of sheaths glabrous; base of inflorescence primary branches glabrous 

4. Inflorescences 4.0-9 cm wide, 12-18 cm long, open; branches 

spreading; plants inland, not coastal S. potosiensis Wipff & Jones 

4. Inflorescences 0.6-1.0 cm wide, 2-8 cm long, contracted; branches 

appressed; plants coastal S. virginicus (L.) Kunth 

Sporobolus potosiensis Wipff & S.D. Jones, sp. nov. (Figs. 1-4) 

antheris 3, 1.2-1.4 mm longis, luteolis. 

Vlants (Fig. 1) perennial, erect, 18-5 1 cm tall with slender rhizomes (Fig. 
2) to 14.5 cm long, 1.3-3.5 mm wide. Leaves 4-5, cauline, subcoriaceous; 
sheaths glabrous, margins ciliate; collar (Fig. 4) abaxially glabrous, yellowish, 
margins conspicuously long ciliate with trichomes to 5-5 mm long; blades 
6.2-24.0 cm long, 0.6-2.5 mm wide, flat, folded or involute; abaxial surface 
glabrous; adaxial surface minutely papillose, antrorsely scaberulous along 
raised veins; margins antrorsely scaberulous; ligules 0.1-0.3 mm long, 
ciliolate membrane, truncate, lacking long trichomes behind the ligule. 
Inflorescences an open panicle, 12-18 cm long, 4.0-9-0 cm wide, glabrous; 
lower branches 3.0-8.0 cm long, nonfloriferous in the proximal 1/3-1/2; 
pedicels 0.2-2.2 mm long, glabrous; spikelets appressed to tertiary branches. 
Spikelets (Fig. 3) 1.6-2.5 mm long, 0.5-1.1 mm wide, glabrous, mottled 
purplish, with one floret; glumes unequal; /im glumes 1.0-1.5 mm long. 

FIG. 1. Photograph of >r., 

veinless, lanceolate; jefo;z^^///;;2^j 1.3-2.1 mm long; conspicuously 1-veined, 
vein green; lanceolate with white-hyaline margins in the distal half, slightly 
shorter than lemma; lemmas 1.5-2.3 mm long; conspicuously 1-veined, vein 
green, ovate-lanceolate, mottled purplish with white-hyaline margins in 
distal half;/;^/e^j 1.6-2.2 mm long, 2-veined, grooved or furrowed between 
veins, similar to lemma texture and color; anthers 3, 1.2-1.4 mm long, 
yellowish. C^ryo/jj^j 0.9-1. mm long, 0.4-0. 5 mmyNxde. Chromosome number 

The specific epithet refers to the State of San Luis Potosi, Mexico; the only 

Distribution: Known only from the Rio Verde River Valley, San Luis 
Potosf, Mexico. 

Typus: MEXICO. San Luis Potosi: 9 km al E de Rio Verde, sobre la carretera a Rayon, 
alt. 1000 m, terrenos aluviales pianos con vegetacion de zacatal halofilo con Prosopis, 
graminea dominance, 16 Sept 1967,/. Rzedoivskt 24790 (holotype: MICH!; isotypes: LL!, 

Additional specimens examined (paratypes): MEXICO. San Luis Potosi: in the valley of 

(TAES); in the valley of the Rio Verde and in the Sier 

San Francisco-Patios-Cardenas-Rayon, 14-15 Sept l95A,Sohm 

del Maiz, 0.7 mi N of Las Tablas (RR crossing in town), alkal 

Samolus, Vig/mra, Flaveria, alt. 1010 m, 22°17'N, 99°52'W, 
Wells (ARIZ, TEX); ± 5 km al SW de Tablas, alt. 1000 m, 2 

(MEXU); Region de Llanos de Angostura municipio de Rio Verde, alt. 900 m, 29 Jul 1980, 
Bravo 14 (MEXU); 0.65 km (0.4 mi) N of the Escuela Primera in Las Tablas, on gravel road 
running between Hwys 70 and 80, alt. 1097 m (3600 ft), 27 Jul 1979, Lane 25386 &J. E. 
Fryxelt (TEX). 


We thank Tony A. Re2nicek (MICH) for his assistance with the Latin 
description and review of the manuscript. We thank Charlotte G. and John 
R. Reader (ARIZ), and an anonymous reviewer for their review and 
suggestions. We also would like to thank the Curators at the following 
herbaria for loaning specimens: GH, MEXU, MICH, MO, TEX/LL, and US. 


Chase, A. 1951. 

5 Man 

2nd ed. U.S.D.A. Misc. Publ. No. 200. United State 

Washington, D.C. 
Clayton, W.D. and S.A. Renvoize. 1986. Genera gramin 

Bull., Addit. Ser. XIIL Her Majesty's Stationery Office 
CoRRELL, D.S. and M.C. Johnston. 1970. Manual of the 

, grasses of the World. Kew 

1 Foun. 

, TX. 

WlPFF AND JONES, Sporobolus 1 69 

Gould, F.W. and R.B. Shaw. 1983. Grass systematics, 2nd ed. Texas A&M University Press, 

College Station, TX. 
Holmgren, P.K., N.H. Holmgren and L. C. Barnett. 1990. Index herbariorum. Part I: The 

herbaria of the world. Regnum Veg. 120. New York Botanical Garden, New York, NY. 
Johnston, I.M. 1943. Plants of Coahuila, eastern Chihuahua, and adjoining Zacatecas and 

Durango, II. J, Arnold Arbor. 24:377-421. 
JuDZiEWicz, E.J. and P.M. Peterson. 1989- Sporobolus temomairemensis (Poaceae: Eragros- 

PoHL, R.W. 1980. Gramineae. In: Flora Costaricensis, Family #1 5, ed. W. Burger. Fieldiana, 

Botany No. 4. Field Museum of Natural History, Chicago, IL. 
Reeder, C.G. 1975. Sporobolus. In: The grasses of Texas, Frank W. Gould. Texas A&M 

University Press, College Station, TX. 


The Department of Botany of the National Museum of Natural History, 
Smithsonian Institution, is pleased to announce the inauguration of a 
Gopher Server that will provide Internet access to databases and documents 
that our staff have created or now manage. The enclosed brochure describes 
these offerings. 

We wish to call your attention in particular to the Type Specimen 
Register of the U.S. National Herbarium, which with over 88,000 
records is the world's largest electronic database of plant types and will 
become an important reference for monographers and flora writers. Also 
please note that our Index to Historical Collections is the most compre- 
hensive account of the collectors represented in the U.S. National Her- 
barium. Neither of these databases were available electronically or inprinted 
form before. 

Our Gopher Server provides electronic access with key word searching to 
several documents that previously were available only as printed matter. 
These include the Checklist of the Plants of the Guianas (Guyana, Surinam, 
French Guiana) (1 992), the Bwlogkal Conservation Newsletter ( 1 98 1 -present) 
and, on behalf of the American Society of Plant Taxonomists, the ASPT 
Newsletter (1987-present). 

We hope you will bring this announcement to the attention of your 
readers. We would also appreciate expressions of support for our efforts to 
make this information available. A strong and favorable response from the 
botanical community will enable us to make more databases and documents 
available in the future. 






So/ahem Weed Science Laboratory 
Stonevtlh, MS 58776, U.S.A. 


Herbarium, Department of Biology 

Valdosta State University 

Valdosta, G A 51698, U.S.A. 

rex htckmllii 
■grostts, C. lo. 
vifolmdes, ha 

f, in the southeastern United States. Addition 
egatus, C. dtfformts, C. flavtcomus, C. lancastre 
. Locality and habitat data are presented for a 

. entrerianus. C. eragrostts, C. iouisianensisy Kyllingabrevifolioides. Dosdeestos,C. entrerianus 
K. hrevifoioides. tienen potencial para llegar a ser una amenaza como malas hierbas para la agri- 
jltura y los cespedes, respectivamente, en el sudeste de los Estados unidos. Se presentan am- 

, pilos//s en Mississippi. Se presentan dates de localidad y habitat de todas las especies citadas. 

While continuing to prepare a synoptical treatment oiCarex, Cyperus, and 
Kyllinga as a contribution to the Flora of Mississippi Project, we have 
examined herbaria specimens and conducted field surveys for species with 
potential to occur within Mississippi. We have also continued assessment of 
population size, distribution, and habitat requirements of recently reported 
species, especially those which have potential to become weedy. This article 
adds to the knowledge of Carex, Cyperus, and Kyllinga which has been 
reported in recent years (Bryson 1984a; Bryson & Jones 1990; Bryson et al. 

1991; Bryson & Carter 1992; Bryson et al. 1992; Carter et al. 1987; Morris 
& Bryson 1 986; Naczi & Bryson 1990). As we previously discussed, tlie flora 
of Mississippi is still poorly known in comparison with several adjacent 
states. Lowe's Plants of Mississippi {1921), although outdated, must continue 
to serve as a base line for the general floristic work in the state. Other 
references that we used as sources of distributional data are Mackenzie 
(1931-1935), Kukenthal(1935-1936),McGivney(1938),Corcoran (1941), 
Horvat (1941), Radford et al. (1964), and Godfrey & Wooten (1979). 

The terminology of physiographic regions or resource areas in Mississippi 
follows Lowe ( 1 92 1 ) as adapted by Morris (1989). Herbarium abbreviations 
follow Holmgren et al. (1990), except ctb, MMNS, and USMH (personal 
herbarium of Charles T. Bryson, Mississippi Museum of Natural Science, 
Jackson, and University of Southern Mississippi, Hattiesburg, respectively). 


Carex bicknellii Britton var. opaca F.J. Herm. was described from three 
collections by Dr. Delzie Demaree from river terraces in Lonoke and Prairie 
counties, Arkansas (Hermann 1972). In his discussion of this puzzling 
variety of C. bicknellii, Hermann (1972) emphasized its large, nearly 
nerveless, and partially translucent perigynia and indicated they were 
similar to perigynia of C. brittoniana Bailey and C. merritt-fernaldii Mack, as 
well as C. bicknellii. However, the perigynia of C. bicknelliivz.\i. opaca are much 
larger (5.5-7 mm long, 4-4.75 mm wide) compared to those of C. merritt- 
fernaldii (4-5 mm long, 2 . 5-3 . 5 wide). Carex bicknellii var. opaca differs from 
the Oklahoma-Texas C. brittoniana in usually having 5-7(9) (rarely only 4) 
spikes per culm; perigynia (2)4-7 nerved (sometimes faintly) over the 
achene ventrally; and staminate and pistillate scales obtuse to long-acumi- 
nate, but with the midrib not excurrent as a scabrous awn (rarely a short 
mucro present). Carex brittoniana has culms with normally (2)3-4(5) spikes; 
perigynia nerveless over the achene ventrally, or nearly so; and staminate 
scales (and sometimes the lowermost pistillate scales) with the midrib 
excurrent as a scabrous awn 0.1-0.9(2.4) mm long. Carex bicknellii var. 
bicknellii is a dry prairie species not known from Mississippi. Carex bicknellii 
var. opaca was found at the same site in the Black Prairie Region of 
Mississippi where C. oklahomensis Mack, was first collected in the state by the 
senior author in 1991 (Bryson et al. 1992). It was associated with C. biishii 
Mack., C. complanata Torr. & Hook., C. glaucodea Tuckerman, C. longii 
Mack., and C. vulpinoidea Michx. The following are data reporting C. 
bicknellii var. opaca from Mississippi for the first time. 

Bryson and Carter, Cyperaceae in Mississippi 173 

Carex fissa Mack. var. fissa is herewith reported new to Mississippi. 
Carexfissa was described by Mackenzie (1931) from specimens collected in 
eastern Oklahoma where it was believed to be endemic. Kolstad (1986) 
reported C. fissa from southeastern Kansas and eastern Oklahoma. Hermann 
(1965) described Carexfissa var. aristata from north central Florida. Jones et 
al. (1990) reported C. fissa new to Texas. The Mississippi collection of C. fissa 
yar. fissa was found in a wet ditch between old hwy US 78 and railroad tracks 
in with C. anmctens Bicknell, C triangularis Boeck., and C. vulpinoidea 
Michx. in the North Central Plateau Region. It is possible that this species 
may have been introduced into Mississippi by highway or rail traffic. 
However, its habitat in Mississippi is similar to that in Arkansas, and it is 
likely that C. fissa is native to Mississippi. It is probable that C. fissa var. 
aristata will be found in southern Mississippi. The following data report C 
fissa vat. fissa new to Mississippi. 

Voucher specimens: U.S.A. MIJ 

Cyperus drummondii Torr. & Hook. [ = C. virensM.ich.-K. var. drummondii 
(Torr. & Hook.) Kukenthal} in North America is restricted to the coastal 
plain and is known from Texas and Louisiana (Denton 1978). Specimens 
have also been seen from Florida and Georgia (Carter, unpublished data). It 
is also known from Nicaragua, Jamaica, the Galapagos Islands, Surinam, and 
Brazil (Denton 1978). Although Denton (1978) considered C. drummondii 
to be a variety of C. virens Michx., we think the morphological differences 
are sufficient for recognition as a species. Cyperus drummondii is generally a 
taller plant with a more compact inflorescence and fewer primary inflores- 
cence bracts, shorter scales, and a greater achene length to scale length ratio. 
At the site reported here, C. drummondii was associated with C. haspan L., C 
ovatus Baldwin, C. polystachyos Rottb., C. strigosus L., C. virens, and Fimbri- 
stylis spp. in the Coastal Pine Meadows Region. The following are data for 
the first collection of C. drummondii from Mississippi. 

Voucher specimens: U.S.A. MISSISSIPPI. Jackson Co.: Moss Point, ditch along W side 
of hwy MS 63 between Frederick Street and Dr. Martin Luther King Drive, 0.5 mi S 
Escatawpa River Bridge, 16 Sep 1993, Carter 11515 (ctb, SWSL, VSC). 

Cyperus elegans L. is known from coastal Texas, Louisiana, and Florida; 
it has not previously been reported from Mississippi. It is related to C, 
oxylepis Nees ex Steud. and can be distinguished from that species by its 
black, obovoid achenes (0.7—1.0 mm broad) and grayish brown, semi- 
translucent scales versus the brown, oblong, ca. 0.5 mm broad achene and 
golden yellow to orangish scales of C. oxylepis (Bryson & Carter 1992). Like 
C. oxylepis, C. elegans has a distinctive and pleasantly aromatic fragrance 

somewhat like that of cured juniper wood, which can often be detected 
several meters away from live plants in the field. The authors have on several 
occasions smelled these species in the field prior to making visual contact. 
Both species are locally abundant in heavily disturbed fill areas along the 
coast in Jackson County, Mississippi, and the populations of C. elegans and 
C. oxylepis consist of more than 2000 plants each and cover an area greater 
than 121 ha (300 acres). Cyperus elegans was found in association with C. 
odoratus L, C. oxylepis, C. entrerianus Boeckler, C. filidnus Vahl, C. flavescens 
L.,C. surinamensis Rottb., and C. virens in the Coastal Pine Meadows Region. 
Collection data for C. elegans in Mississippi follow. 

Collection data. U.S.A. MISSISSIPPI. Harrison Co.: Orange Grove Community Center, 
0.3 mi N jet. hwysI-10andUS49;WofUS49,l6Oct 1993, Br^^oK 13160 ^«^Ctfner(ctb, 
SWSL, VSC). Jackson Co.: Pascagoula, vicinity of Bayou Casotte, S jet. of Louise and 
Washington Streets, 9 Aug 1993, Bryson 12595 (ctb, IBE, MICH, VDB, VSC); Carier 
11339 (VSC, others to be distributed). 

Cyperus entrerianus Boeck. is an apparent introduction from South 
America or Mexico (Carter 1990) and was reported from 21 counties in 
Georgia, Florida, Alabama, Louisiana, and Texas (Carter 1990; Carter & 
Jones 1991). Although C, entrerianus is not recognized m some floras, it is 
quite distinct from other taxa in the Cyperus section Luzuloidei in temperate 
North America, and a comprehensive discussion and key was provided for 
this section by Carter (1990). This perennial is a copious producer of seeds 
and possesses a hard, stout, deeply set rhizome. It seems to be a more 
aggressive weed than other members oiCyperus section Luzuloidei that occur 
in Mississippi {t.g.,C. acuminatus Torr. & Hook., C. drummondii, C. eragrostis 
Lam., C. pseudovegetus Steud., C. surinamensis, and C. virens). Cyperus entreri- 
anus is often locally abundant in eastern Texas and southern Louisiana where 
it seems to be associated with rice production, and it has been found in a rice 
field in Paraguay (Carter 1990). Cyperus entrerianus was found growing in the 
Coastal Pine Meadows Region with C. elegans and its associates listed in the 
preceding discussion. The following are data for the first collections of C. 
entrerianus in Mississippi. 

Voucher specimens: U.S.A. MISSISSIPPI. Jackson Co.: Pascagoula, vicinity of Bayou 
Casotte, S jet. of Louise and Washington streets, 9 Aug 1993, Bryson 12598 (ctb); 31 Aug 
1993, Bryson 12821 and Newton (BRIT/SMU, ctb, DSC, FLAS, FSU, GA, IBE, KNK, 

Cyperus eragrostis Lam. is native to California, Oregon, and Washing- 
ton in North America; Bolivia, Peru, and Surinam south to Argentina, 
Brazil, Chile, and Uruguay in South America; and Easter and Juan Fernan- 
dez Islands in the Pacific (Denton 1978). This species has not previously 

Bryson and Carter, Cyperaceae in Mississippi 1 7 5 

been reported from Mississippi although two specimens collected by Ken 
Rogers (cited below) were collected in Forrest and Hinds counties in 1971 
and 1981, respectively, and determined by R. Krai (VDB) and R. Carter. In 
Rankin County, collections were made along shallow narrow ditches. These 
sites are across the Pearl River and within five miles of the earlier Hinds 
County site of Ken Rogers. The IMississippi plants seem to be intermediate 
in size between typical C. eragrostis var. eragrostis and the diminutive form C. 
eragrostis var. compactus (Desv.) Kuk. Denton (1978) treated C. eragrostis var. 
compactus as a synonym of C. eragrostis because the diminutive form occurs 
throughout the range of the species. When transplanted into pots and grown 
under controlled conditions (i.e., with regular watering and fertilizer) in the 
greenhouse at the USDA, ARS, Jamie Whitten Delta States Research 
Center, Stoneville, MS, individuals of C. eragrostis from the Rankin County 
collection produced new stems that were longer than those observed in the 
field and that were typical of Cyperus eragrostis var. eragrostis. Thus, it appears 
periodic mowing of ditches and roadsides apparently caused plants to be 
shorter than typical for the species in the Rankin County, Mississippi 
population observed by the senior author. In Rankin County, C. eragrostis W3.s 
associated with Carex longii, Cyperus haspan, C. odoratus., C. pseudovegetus , C. 
strigosus, C. virens, and Kyllmga brevifolia Rottb. { = Cyperus hrevifolius (Rottb.) 
Hassk.}. Each of the collections were made in the Longleaf Pine Belt Region. 
The following data are the first records C. eragrostis from Mississippi. 

Voucher specimens: U.S.A. MISSISSIPPI. Forrest Co.: Hattiesburg, drainage ditch at 
Kemper Park, 16 Jul \91\,Rogen 6708 (VDB). Hinds Co.: Jackson, Riverside Park, 8 Jun 
1981, i?<?gm46973 (VDB). Rankin Co.: Flowood, 0.3 mi Eof Pearl River;Sofhwy MS 25, 
7 Sep 1993, Bryson 12966 (ctb, VSC); Bryson 12975 (ctb, SWSL, VSC); Flowood, 0.7 mi E 
Pearl River; S of hwy MS 25, 14 Oct 1993, Bryson 13128 and Bryson (ctb, SWSL, VSC). 

Cyperus louisianensis Thieret was previously reported from only two 
sites, including the type locality, in Tangipahoa Parish, Louisiana (Thieret 
1977). Its lenticular achene and two-branched style place it in subgenus 
Pycreus, and it appears to be closely related to the North American species, 
C. diandrus Torr. (Thieret 1977). The scales of C. louisianensis are similar to 
those of C, diandrus, but C. louisianensis is distinguished by three stamens and 
styles that are divided less than halfway to the base (Thieret 1977). Cyperus 
louisianensis is also closely related to the widespread old world species, C. 
sanguinolentus Vahl (Thieret 1977). Our observations indicate that C, 
louisianensis is typically found in disturbed habitats, such as road ditches and 
margins of artificial ponds, where it often forms dense, nearly monotypic, 
stands. Also, it is found in association with introduced weeds, e.g. C. pilosus 
Vahl (Bryson & Carter 1992) and Sacciolepts indtca (L.) Chase (Bryson & 
Lockley 1 993), in the Coastal Pine Meadows Region. We have also observed 

that C. louisianensis flowers and fruits from late August until frost and that 
it is much more easily detected late in the season when its distinctive reddish 
scale pigmentation has developed completely. Thus, this may explain why 
it has been overlooked previously in Mississippi. Cyperm louisianensis has 
been listed as a candidate for protection (U.S. Fish and Wildlife Service 
1993). Additional field and herbarium studies by the authors are currently 
in progress to determine the status of this taxon. The following data are the 
first records of C. louisianensis from Mississippi. 

Collection data. U.S.A. MISSISSIPPI. Hancock Co.: W side of hwy MS 43, 0.44 mi N 
of jet. hwys US 90 and MS 43 in Waveland, 16 Sep 1993, Carter 11342 (VSC, SWSL, others 
to be distributed); I6 Oct 1993, Bryson 13166 and Carter 1 154^ (ctb, SWSL, VSC, others 
to be distributed); W of Mill Creek, between Mill Creek and Indian Ridge Road, S of MS 
43, ca. 5 mi E jet. hwys MS 43 and 1-59 in Picayune, 18 Oct 1993, Bryson 13265 and Carter 

11567 (ctb, SWSL, VSC, others to be distributed); 9.0 mi NW of jet. hwys 43 and 603 m 

11568 and Bryson (VSC, others to be distributed); N of Kiln, 0.6 mi S of jet. hwys MS 43 
and MS 603, byW side of hwy MS 43, 18 Oct 1993,Brj.;w; 13267 and Carter 11569 {cth, 
SWSL, VSC, others to be distributed); Kiln, beside Shitalo Baptist Church and across hwy 
MS 43 from Kiln Post Office, by W side of hwy MS 43, 18 Oct 1 993, Brjw 7 3268 <7;2^C^r?^r 
U570 (ctb, SWSL, VSC, others to be distributed); SE jet. hwys MS 43 and I-IO, 18 Oct 
1993, Br^Jo«i3277^Wc:^r;fr/ 757/ (ctb, SWSL, VSC, others to bedistributed). Harrison 
Co.: Orange Grove Community Center, 0.3 mi N jet. hwys I- 10 and US 49; W of US 49, 
16 Oct 1993, Bryson 13164 and Carter 11544 (ctb, SWSL, VSC, others to be distributed); 
Orange Grove, 1. 1 mi S jet. hwy US 49 and O'Neal Road, 18 Oct 1993, Bryson 13276 (ctb, 
SWSL, VSC, others to be distributed); N Gulfport, 0.3 mi W of jet. of Harrison Drive and 
34th Avenue, ditch along Harrison Drive, 18 Oct 1993, Carter 11574 (VSC, others to be 
distributed); 3.32 mi W of jet. of Popps Ferry Road and D'Iberville Boulevard ( = hwy MS 
67), along Popps Ferry Road, ea 50 m W of jet. with Camp Four Jacks Road, 18 Oct 1993, 
Carter 11577 (VSC, others to be distributed); NW jet. hwys I-IO and US 49, 18 Oct 1993, 
Bryson 13279 (ctb, SWSL, VSC, others to be distributed). Jackson Co.: Paseagoula, just SE 
jet. of Washington and Louise Streets, vie. Bayou Cassette, 1 6 Sep 1991 , Bryson 11032 and 
be distributed); 16 Sep 1993, Carter 11337 (VSC, others to be distributed); St. Martin, 0.2 
mi N of jet. of Old Fort Bayou Road and Rosefarm Road, along Rosefarm Road S of creek, 

W of jet. of Old Fort Bayou Road and Yellow Jacket Boulevard, between Lancaster Road and 
Mayfair Road, 18 Oct 1993, Carter 11580 (VSC, others to be distributed); vicinity of St. 
Martin, 0.19 mi E of jet. of Fort Bayou Road and Yellow Jacket Drive, by Old Fort Bayou 
Road, 18 Oct 1993, Carter 11581 (VSC, others to be distributed). Pearl River Co.: 

Oct 1993, Bryson 13222 and Carter 11565 (ctb, SWSL, VSC, others to be distributed); 
Picayune, 0.5 mi W jet. hwys 1-59 and MS 43; N of MS 43, 18 Oct 1993, Bryson 13257 and 
Carter 11562 (ctb, SWSL, VSC, others to be distributed). 

Kyllinga brevifolioides (Thieret & Delahoussaye) G.C. Tucker { = C. 
brevifolioides Thieret & Delahoussaye) (Tucker 1984) was cited from Con- 
necticut, North Carolina, Pennsylvania, and Virginia in North America by 

Bryson and Carter, Cyperaceae in Mississippi 177 

Delahoussaye and Thieret (1967) in the original description. Subsequently, 
it has been reported from Maryland (Sipple 1978; Naczi et al. 1986), 
Tennessee (Krai 1981; Webb et al. 1981), Alabama and Georgia (Webb et 
al. 1981), New Jersey (Snyder 1983, 1984), Delaware (Naczi 1984; Naczi 
et al. 1986), and Arkansas (Sundell & Thomas 1988). Like K. brevifolia, K. 
hrevifolioides is a rhizomatous perennial; however, it can be separated from K. 
brevifolia by its smooth scale keel and 2-3 stamens versus the denticulate 
scale keel and a single stamen of K brevifolia. Like K. brevifolia and K. odorata, 
K. brevifolioides is weedy and was probably introduced from Asia (Ferren and 
Schuyler 1 980; Webb & Dennis 1981). Kyllinga brevifolia and K, brevifolioides 
have been observed by the authors to be weeds in periodically wet or 
frequently irrigated areas of lawns, roadsides, ditches, cemeteries, golf 
courses, and flower beds, often associated with the following sedges: Carex 
longii, Cyperus polystachyos, C. pseudovegetus , C. strigosus, Eleocharis obtma 
(WiUd.) Schult., and Kyllinga pumila Michx. The collections reported herein 
are from the Loess Bluffs and Tennessee River Hills Regions. The following 
are data for the first collections oi K. brevifolioides from Mississippi. 

Voucher specimens: U.S.A. MISSISSIPPI. Alcorn Co.: Corinch, S of jet. hwy US 72 and 
Cass Street, 29 Sep 1993, Bryson 13082 (ctb, FSU, IBE, KNK, NYS, SWSL, VDB, VSC); 
Corinth, SE of jet. hwy US 72 and Harper Road, 29 Sep 1995, Bryson 13094 (ctb, IBE, NLU, 
SWSL, VDB, VSC). De Soto Co.: Olive Branch, 0.3 mi W of jet. hwys US 78 (old) and MS 
302 ( = Goodman Road); N of Goodman Road, 7 Oct 1995, Bryson 13102 and Bryson (BRIT/ 
VSC); ca. 0.8 mi N jet of Goodman Road and Getwell Road; W of Getwell Road, 13 Oct 

Cyperus aggregatus (Willd.) Endl. {previously known as C. huarmensis 
(H.B.K.) M.C. Johnst., C cayennensis (Lam.) Britton, and C. flavus (Vahl) 
Nees; see Tucker (1985)} is reported from Florida (Krai 1966; Clewell 
1985),Louisiana(Horvat 1941), and Texas (Correll&Johnston 1970). Since 
It was first discovered in Stone County, Mississippi, in 1991 (Bryson & 
Carter 1992), C. aggregatus has been observed to be an aggressive weed of 
open sandy hilltops, roadsides, poorly kept lawns, and disturbed vacant lots 

distribution of populations in Mississippi suggest that C. aggregatus, if not 
a native, was introduced many years ago into the Coastal Pine Meadows and 
Longleaf Pine Beit Regions of Mississippi. The following are additional 
records of this weedy species, including two new county records. 

Voucher specimens: U.S.A. MISSISSIPPI. Forrest Co.: Fruitland Park, E of hwy US 49, 
9 Aug 1993, Bryson 12580 (ctb, IBE, SWSL, VSC). Harrison Co.: 0.2 mi N of Howison, 
E of hwy US 49, 9 Aug 1993, Bryson 12381 (ctb, IBE, NYS, VSC); Gulfport, vacant lot W 

Cyperus difformis L. ranks as one of the world's worst weeds (Holm et 
al. 1991) and occurs from Europe, Asia, Central America, North America 
(Lipscomb 1980), Australia, the Pacific Islands (Kukenthal. 1935), Mexico 
(McGivney 1938), and South America (S. McDaniel pers. comm.). It is a 

period (as little as 4 to 6 weeks from seed to seed) and high reproductive 
potential (Holm et al. 1991). This species seems to be spreading in the 
United States especially along major waterways, through introduction at 
ports-of-entry, and in rice production in California. In the United States C. 
difformis has been known from Virginia since 1 934 and is a troublesome weed 
of rice in California (Bryson 1984b). It is also known from Alabama (Krai 
1973), Florida (Burkhalter 1985), Louisiana (Thieret 1964), Mississippi 
(Bryson & Carter 1992), Nebraska (Lemaire 1970), North Carolina (Tyndall 
1983), Pennsylvania (Smith 1986), Tennessee (Webb & Dennis 1981), and 
Texas (Carr 1988). During field work in 1993, new populations were 
discovered, and previously known populations (Bryson & Carter 1992) were 
observed. Population size, in area covered and number of individuals, had 
increased by 2- to 400-fold since 1991. The following are data for recently 
discovered populations of C. difformis in Mississippi from the Coastal Pine 
Meadows and Longleaf Pine Belt Regions. 

Voucher specimens: U.S.A. MISSISSIPPI. Forrest Co.: Hattiesburg, Alcorn Street 
between James and Tipton streets, beside Hattiesburg Water Works, 16 Oct 1993, Bryson 
13148, Carter anci Rosso (BRIT/SMU, ctb, FSU, IBE, KNK, MISS, MMNS, MO, NLU, 
NYS, SWSL, TAES, USMH, VDB, VSC). Hancock Co.: Kiln, wet ditch W of hwy MS 43, 
6 Aug 1992, Bryson 11941 (ctb, IBE, VSC). Harrison Co.: Gulfport, wet ditch between 
Seaway Road and RR; S of hwy I-IO, 6 Aug 1992, Bryson 11993 (ctb, IBE, VSC); Orange 
Grove, 0.3 mi N jet. hwys I-IO and US 49; W of US 49, 31 Aug 1993, Bryson 12805 and 
Newton (ctb). Jackson Co.: Moss Point, SE jet. hwys I- 10 and MS 63, 31 Aug 1993, Bryson 
12818 and Newton (ctb); Moss Point, NE of jet. hwys US 90 and MS 63, 1 Sep 1993, Bryson 
12833 and Newton (ctb, IBE, NLU, VDB, VSC); Pascagoula, Bayou Casotte area, adjacent 
to City Animal Shelter just S of S end of Louise Street, 16 Sep 1993, Carter 11338 (VSC, 

:hx. (=C. albomarginatus Martius & Schrad. ex 
Nees) was previously known from only Lafayette and Tishomingo counties, 
both in northeastern Mississippi in the North Central Plateau and Tennessee 
River Hills Regions. Cyperus flavicomus was found growing on wet soil often 
in shallow standing water in association with Carex longii, Cyperus difformis, 
C. esculmtus L., C, iria L., C. haspan, C. odoratus, C. pilosus, C. polysfachyos, C. 
strigosus, C. surinamensis, C. virens, Eleocharis obtusa, E. tuberculosa (Michx.) 
Roem. & Schult., Fimbristylis autumnalis (L.) Roem. and Schult., F. miliacea 

Bryson and Carter, Cyperaceae in Mississippi 179 

(L.) Vahl, Kyllinga hrevifolia, and Rhynchospora cornkulata (Lam.) Gray. Based 
on our examination of herbarium specimens and field surveys, C . flavkomus 
seems to occur sporadically throughout much of the southeastern United 
States. The following are additional county records from the Coastal Pine 
Meadows Region of southern Mississippi. 

Voucher specimens: U.S.A. MISSISSIPPI. Harrison Co.: Orange Grove, 0.3 mi N jet. 
iiwys I- 10 and US 49; WofUS 49, 31 Aug 1993,6rj'Jo« i2Si 5 «WNf«./o«(ctb,DSC,IBE, 
MISS, MMNS, MO, NLU, SWSL, USMH, VDB, VSC); Bryson 13162 and Carter {czh, VSC). 
Jackson Co.: Moss Point, SE of jet. hwys I- 10 and MS 63, 31 Aug 1995, Bryson 12816 and 

Cyperus lancastrensis Porter in Gray was first reported from Mississippi 
by Morris ( 1 988) in the North Central Plateau Region. Subsequently, it was 
reported from Lee and Tishomingo counties in the Tennessee River Hills 
Region (Bryson & Carter 1 992). Following are data from an additional three 
counties all from northern Mississippi in the North Central Plateau and 
Tennessee River Hills Region. Cyperus lancastrensis was found growing in 
association with C. echinatus (L.) Wood and C. strigosus in small damp 
depressions on open hillsides. 

Voucher specimens: U.S.A. MISSISSIPPI. Itawamba Co.: 1.6 mi N Dorsey SW of jet. 
of hwy US 78 and Fawn Grove exit, 26 Aug 1993, Bryson 12784 (ctb, VSC). Marshall Co.: 
N of Galena, jet. of old and new hwys MS 7, 24 Aug 1993, Bryson 12731 (ctb, IBE, VDB, 
VSC). Tate Co.: Thyatira, S of hwy MS 4, 7 Oct 1993 Bryson 13114 and Bryson (ctb, VSC). 

Cyperus ovatus Baldwin { = C. pollardii Britton in Small) occurs through- 
out Florida and northward in the outer Atlantic coastal plain into North 
Carolina and westward along the Gulf Coast into Louisiana (Carter, in prep.). 
It is related to C, retrorsus and is discussed previously from Mississippi 
(Bryson & Carter 1992). At the time of our previous paper, we were unable 
to locate C. ovatus in Hancock County despite field surveys in suitable 
habitats (e.g., pond margins, edges of salt marshes, banks of black water 
streams, and bogs and ditches in pineland savannas). Following are data for 
additional recent collections of C ovatus from Mississippi in the Coastal Pine 
Meadows Region. 

Voucher specimens: U.S.A. MISSISSIPPI. Hancock Co.: Port Bienville Waterfront 
Industrial Park, 17 Oct 1993, Bryson 13182 andCarter 11333 (ctb); Kiln, W of hwy MS 43; 
across hwy MS 43 from Kiln Post Office, beside Shifalo Baptist Church; ca. 2.0 mi S jet. hwys 
MS 43 and 603, 18 Oct 1993, Bryson 13269 andCarter (ctb). 

Cyperus pilosus Vahl, a weed of tropical and subtropical regions of Asia, 
Australia, and West Africa (Kukenthal 1935, McGivney 1938), was appar- 
ently introduced into North America from the old world and was first 
reported in the United States from Tangipahoa Parish, Louisiana, by O'Neill 
(1938). Subsequently, C. pilosus was reported from Florida by Burkhalter 

(1985), relocated in Louisiana by Carter (Bryson & Carter 1992), and 
discovered in Mississippi by Bryson & Carter (1992). In Mississippi, C. 
pilosus was found most frequently growing in disturbed soil at the edge of 
standing water in ditches, along slow moving streams, and around ponds 
often in association with many other Cyperus species including: C. difformis. 
C. ma, C. louisianensis, C. odoratus, C. strigosm, and C. virens. The following 
data report C. pilosus from additional counties in Mississippi in the Coastal 
Pine Meadows Region. 

Voucher specimens: U.S.A. MISSISSIPPI. Hancock Co.: SE of Napolean, in ditch E of" 
jet. hwys I-IO and MS 607; N of I-IO, 10 Aug 199.^, Bryson 12606 (ctb, IBE, MO, NLU, 
SWSL, VDB, VSC); N of Waveland, W of hwy MS 43, 0.44 mi N of jet. hwys US 90 and 
43, 16 Sep 1 99^ Carnr 11346 (VSC:). Harrison Co.: Gulfport, ditch between Seaway Road 
and RR; S of U 1 0, 6 Aug 1 992, Bry.wtj 1 1 996 (ctb, IBE, MICH, NLU, TENN, VDB, VSC); 
Gulfport, ditch along Industrial Road, 9 Aug 1993, Bryson 12582 andLockley (ctb); Bryson 
12584 (ctb, IBE, SWSL, VSC); Bryson 12625 (ctb, IBE, VDB, VSC); Orange Grove 
Community Center, 0.3 mi N )Ct. hwys I- 1 and US 49; W of US 49, 31 Aug 1993, Bryson 
12813 and Newton (ctb, VSC); Bryson 1 3 158 and Carter (ctb, VSC). Pearl River Co.: Ca. 1 


Thanks are expressed to N.B. Bryson, T.C. Lockley, and T.E. Newton for 
help with field work and A.A. Reznicek (MICH) who confirmed the 
determination oiCarex bickmllii var. opaca and C. fissa y2.t.fissa, J. W. Thieret 
(KNK) who confirmed the determination of the 1991 Cyperus louisianensis 
collection, and S.D. Jones (TAES), R.F.C. Naczi (KNK), and A.A. Reznicek 
who reviewed the manuscript. Financial support of the junior author's field 
work was provided by a U.S. Fish and Wildlife Service status survey contract 
on Cyperus louisianensis. 


a sedge. Soui 

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1986. New records of some rare DelMarVa sedges. Bartonia 52:49-57. 
Naczi, R.F.C. and C.T. Bryson. 1990. Noteworthy records of Ctfrex(Cyperaceae) from the 

southeastern United States. Bartonia 56:49-58. 
O'Neill, H.T. 1938. Cyperus pdosus Vahl in the United States. Rhodora 40:74. 
Radford, A.E., H.E. Ahles and C.R. Bell. 1964. Manual of the vascular flora of the 

Carolinas. The University of North Carolina Press, Chapel Hill. p. 203. 
SiPPLE, W.S. 1978. An atlas of vascular plant species distribution maps for tidewater 

Maryland. Wetland Publ. No. 1. Maryland Dept. Nat. Res., Annapolis. 
Smith, T.L. 1986. News and notes: eastern Pennsylvania natural diversity inventory 1985 

field highlights. Bartonia 52:83-84. 
Snyder, D.B. 1983. Rare New Jersey grasses and sedges. Bartonia 49:71-72. 

^cords oi Cyperus (Cyperaceae) in Arkansas. 

SuNDELL, E. anc 

\ R.D. Thomas. 

1988. Four ne 

Sida 13:259 



1964. More adc 

litions to the 

1977. Cyperus 


Louisiana. L 

ouisiana Acad. 

Sci. 40:23-26 

Tucker, G.C. 

1984. Arevisio 

.n of the genu 

Central Am< 

;nca. Rhodora 1 
1985. The ro: 

36: 507-538. 
rrecr name fo 

Kylltnga Rottb. (Cyperaceae) in Mexic( 

Cyperus cayennensis (C, flavus), Cypera 

Cyperus difformis L. (Cyperus) in the southea 

Plant taxa for listing as endangered or threat 

Webb, D.H. and W.M. Dennis. 1981 . Additions to the flora of Tennessee. Sida 9: 184- 

Webb, D.H., W.M. Dennis and T.S. Patrick. 1981. Distribution and naturalizati^ 

Cyperus brevifolioides (Cyperaceae) in eastern United States. Sida 9:188-190. 


hw. Naturalis 

t 30:607-608. 


LL, R.W. 198^ 

>. Distribution of 

:ed States. Cas 


U.S. Fi 

iFE Service. 1993. 



The Tucker Center 

College of Forestry 
Stephen F. Austin State University 
Nacogdoches, TX 7^962, U.S.A. 

(Rupr.) Rupr. genhemis S. Y. Li & K.T. Adair and Pmus sylvestri 
& S. Y. Li) S.Y. Lj & K.T. Adair and Penthorum sedoides L. subsp. t 

Larixgmelini (Rupr.) Rupr. subsp./var. genhemis S.Y. Li & K.T. Adair y Pinus sylvestns L. var. 
manguiensts S.Y. Li & K.T. Adair. Tambien se presentan dos combinaciones nuevas Polem- 
omim hingganicum (RH. Huang & S.Y. Li) S.Y. Li & K.T. Adair y Penthorum sedoides L. subsp. 
chinense (Pursh) S.Y. Li & K.T. Adair. 

This article describes two new taxa and two new combinations from China. 
The study is largely based on collections Shiyou Li made in a four-year forest 
resource investigation in Northeast China during 1984-1987. The speci- 
mens are mainly deposited in the herbarium of Northeast Forestry Univer- 
sity (NEFI) China. 

Larix gmelini (Rupr.) Rupr. var. genhensis S.Y. Li & K. T. Adair, var. nov. 
(Fig. 1) 

Larixgmelini (Rupr.) Rupr. var. genhensis (Genhen larch) is distinguished 
from var. gmelini (Dahurian larch) by its long, slender and pendulous 
branches. The first-year branches of var. genhensis are about two times longer 
than those of var. gmelini. The tree reaches 20 meters tall and 50 cm in 
diameter in favorable conditions. 
Type: CHINA. Neimonggu: Genhe, 4 Jul 1986, S.Y. Li 861-643 (holotype; NEFI!). 

Larix gmelini (Rupr.) Rupr. var. genhensis is sparsely distributed in Genhe, 
Da Xingan Ling, Neimonggu, China (Fig. 3). It usually grows in wetlands 
or occasionally in the Larix gmelini (Rupr.) Rupr. var. gmelini forest on lower 

Pinus sylvestris L. var. manguiensis S.Y. Li & K. T. Adair, var. nov. (Fig. 2) 
Differt a Pinus sylvestris L. var. sylvestris, var. mongolica Litvm., var. sylvestrifomis (Takenou- 

chi) Cheng et CD. Chu foliis 0.5-4.0 cm longis. 

Pinus sylvestris L. var. manguiensis is one of the shortest leaf pines in the 

world. This variety can be distinguished from other varieties of Pinus 

sylvestris by its extremely short and slightly or non-twisted needles (Table 1). 



; Mangui, 26jun 1986, 5. 'K Li 
ined: CHINA. Neimonggu: Jir 

. manguiensis (short-leaf scotch pine) occurs in pure 
stands on dry rocky slopes (400-800 m at elevation) in Mangui and Jinhe, 
Da Xingan Ling, Neimonggu, China (Fig. 3). Pinus sylvestris L. var. 
manguiensis predominates the canopy and is occasionally associated with 
Larix gmelini. The regeneration seedlings under canopy are common, 

mucronulatum Turcz., Spiraea sertcea Turcz., Rosa aciculans Lindl., Vacctnium 
vitis-idaea L. , and Ruhus sp. Main associated herbaceous plants include: Carex 
spp., Calamagrostis sp., Iris ensata Thunb., Pyrola incanata Fisch., Artemisia 
sp., Convallaria majalis L., Pragaria orientalis Lozinsk., and Sanguisorba 
officinalis L. 

The new pine is a beautiful large tree with a short crown and a straight 
reddish-brown trunk. It is usually 20-25 m tall and 50 cm in diameter. It 
is adapted to an extremely continental climate. It has withstood tempera- 
tures of -47°C. The new pine should be managed for timber. 


urn (P.H. Hu 

Pinus sylvestris L. var. manguiensis 
Larix gmelini (Rupr.) Rupr. var. genhensis 

. Bot. Res. 5:151- 

Polemonium htnggankum (Xingan Jacobs-ladder) was named by P.H. 
Huang and S.Y. Li in 1985 and treated as a subspecies of the circumboreal 
P. boreale. Recently, we reexamined this taxon with its closely related taxa. 
The data (Table 2) indicate that the taxon should be treated as a separate 

Li and Adair, Chinese plar 
TABLE 1. Gradient Analysis of 

760 Li 861-604 

NEFI 630082 3.1-7. 
NEFI4001 3.4-7. 

■ sylvesmfc 

Species because it can be distinguished by densely glandular pubescence in 
parts of the plant, large plant size, mostly stem leaves, a long pedicel, short 
corolla, and exposed stamens and stigma. Polemonium htnggankum is similar 
to Polemonium pulchellum in Siberia, but differs from the latter with large size, 
mostly stem leaves, campanulate corolla, and exposed stamens and stigma. 

Penthorum sedoides L. 

Penthorum is a perennial herbaceous genus with a disjunct distribution be- 
tween eastern Asia and eastern North America. Although it is a small genus, 
the taxonomic treatment of species is in dispute. Gray (1846) once doubted 
that the Asian plants (P. chinense and P. burnt le) would be the same as the 
eastern North American R sedoides. Maximowicz (1883) treated P chinense as 
a variety of P. sedoides, and considered P humile as possibly based on abnormal 
specimens. However, in this century, most scholars tend to recognize two 
species P sedoides and P chinense in the genus. Li (1952) clearly stated that P 
chinense differs from P. sedoides in that stems are usually simple or few- 
branched, narrow leaves, has fewer flowered more corybose-like inflores- 
cences and white flowers. In P. sedoides, as Li mentioned, the stems are usually 
many-branched, the leaves broader, the inflorescences more paniculate and 
the flowers greenish. Li's opinion is widely accepted by many scholars. 

The natural range of P. chinense is the northeastern Asia, including Far East 
Russia, Northeast and North China, Korea, and Japan. We exammed all 
available Chinese specimens o( Penthorum and related literature and collected 

hundreds of accounts of specimens during 1985-1987. In 1989, we exam- 
ined the American P. sedoides in both herbarium and field. Finally, we found 
the Asian and American plants to be almost the same in all characteristics 
in nature, with the exception of leaf shape. However, descriptions are often 
individual in flower color. Asian plant flowers have been observed and 
recorded as white, yellow white, and light green; American ones have been 
described as green or yellow-green in much of the literature (e.g. Correll and 
Johnston 1979). Some field notes of specimens are also different: e.g. 
McCrary 1 044 (Texas, 20 Jul 1 984) recorded the flowers as yellow-green, and 
Nixo?? 15992 (Texas, 10 Sep 1987) recorded flowers as white. According to 
our field observations in the last few years, the corollas of Chinese and 
American plants are both white in nature, but they are often inconspicuous 
and fall early while other parts of flowers are all green in color. In most cases, 
the white sepals do not remain or at most are restricted to the upper flowers 
in the inflorescences so that many observers recorded the flowers as green. 
Thus, flower color is not a character on which to distinguish between Asian 
and American plants. 

Further, according to our observations, some Asian plants have simple or 
few-branched stems as Li (1952) described. Others have many-branched 
stems especially in dry and disturbed sites. Therefore, it seems there is no 
difference in stems between Asian and American populations. Also, Li's 
descriptions on the fewer flowered more corybose-like inflorescences (Asian 
plants) and more paniculate (American plants), seem a result of partial 
observations. According to our specimen examinations, there is no distinct 
difference in inflorescences between Asian and American taxa. In addition, 
anatomical data (Haskins and Hayden 1987) failed to show any significant 
difference between these two taxa. 

Actually, the only morphological difference between Asian and American 
taxa is the former has narrow and long willow-like leaves, while the latter 
has wide and shorter glabrous leaves. Geographically, they occupy similar 
habitats on separate continents. Thus, we treat the Asian taxon as a 
geographic subspecies of the American species. As a result, Penthorum 
becomes one of five monospecific disjunct genera in eastern Asia and North 

Penthorum sedoides L. subsp. sedoides 

Distribution. Eastern United States. 

Representative specimens examined. UNITED STATES. Arkansas: D. Demaree 34302 
(ASTC). Iowa; H.B. Wey land 406 (ASTC). Louisiana: C. Smith 1853 (ASTC). Massachu- 
setts: H.E. Ahles 90326 (ASTC). Texas: L.E. Brown 3132 (ASTC); M. McCrary 1044 
(ASTC); E.S. Nixon 7649, 10717, 13992 (ASTC); E.S. Nixon andC. Burandt 7078 (ASTC); 
E.S. Nixon and M. McCrary 3591 (ASTC); E.S. Nixon andJ.R. Ward 10863. 12390, 12422, 
12434, 12547, 12922, 13043, 13136 (ASTC); C. Waters 174 (ASTC). 

190 SiDAl6(l) 1994 

Penthorum sedoides L. subsp. chinense (Pursh) S.Y. Li & K.T. Adair, 

comb, et Stat. nov. Basionym: Penthorum chmensePmsh,m. Amer. Sept. 323. 1811; 

Koma-rov, Fi. Manshur. 2:403. 1903; Kitagawa, Lin. Fl. Manshur. 246. 1939; Liou, 

Key PI. NE China 122. 1959- Penthorum sedoides L. var. chtnensts (Pursh) Maxim., 

Melanges Biol. Bull. Phys. -Math. Acad. Imp. Sci. Saint-Petersbourg. 11:774. 1883. 

Distribution. Korea, Japan, eastern China southward to Indochina. 

Specimens examined. CHINA. Heilongjiang: S.L. Dong 11040 (NEFI); S. Y. Li 125, 84- 

103 (NEFI), S.Q. Nte 964 (NEFI); Y.L. Zhang 1926 (IFP). Jilin: S.Y U 10043 (NEFI). 

Liaoning: S.Y Lt 833007 (NEFI). 

We are grateful to Li-Guo Fu (Chinese Academy of Sciences, Beijing, 
China), Barney L. Lipscomb (BRIT), Shao-Quan Nie (NEFI), and Elray S. 
Nixon and Barrel L. McDonald (ASTC) for reviewing the manuscript. 
Special thanks are due to Puhua Huang and Fucheng Shi (NEFI), Qingsheng 
Meng (Heilongjiang Institute of Traditional Chinese Medicine, Harbin, 
China), Shulan Yin (Ministry of Forestry, Beijing, China), Jing Zhan 
(Mangui Forestry Bureau, Neimonggu, China), and Y. Wang (Genhe 
Forestry Bureau, Neimonggu, China) for aiding field investigations in Da 
Xingan Ling during 1984-1987. 

CORRELL, D.S. and M.C.Johnston. 1979. Manual of the vascular plants of Texas 

Gray, A. 1946. Analogy between the flora of Japan and that of the United Stat 
Arts Sci. II. 2:135-136. (Reprinted in Graham 1972, and Stuckey 1978). 

Haskins, M.L. and W.J. Hayden. 1987. Anatomy and affinities aiPenthorum. A 

!/;.«L. from the Great 

idS.Y.Li. 1985. A 


les ofPolemo 



China. Bull. Bot. 

Res. 5:151- 



PA, m'. 

1939. Lineament 


..1903. Flora of A' 

lanshuria II. 


Li, H.L. 


iships betw. 


. Ame 

r. Philos. Soc. 42: 


Liu SE 

• (ed.) 


us in Northeast China. 


.1965. Flora of Japan. J 




^,N. 1959. Circumpolar Arctic flora 

.. Clarenden 

Welsh, S.L. H 

)74. Anderson's flc 

,ra of Alaska 




208 Mueller Laboratory 

Pennsylvania State University 

University Park, PA 16802, U.S.A. 


Department of Botany 

University of Maryland 

College Park, MD 20742, U.S.A. 

=rous rootstocks occur in Texas: Anemone carolinia 
osa Kydhetg, A. edwardsiana Tharp, and a new speci 
■mic mainly to the Edwards Plateau in west-cent 

Texas cinco espec 


, de Aner, 

none • 

con nzomas tuberc 

3SOs: Ane. 


•ana Walter, 




Tharp, A. tuberosa 

■ Rydber^ 


.- Keener &E 


:on endei 




The genus Anemone'L. consists ofca. 1 50 species of perennial herbs featuring 
one or more radical leaves, involucrate peduncles, petaloid sepals, and achene 
fruits. In a recent preliminary reclassification based on restriction site varia- 
tion and morphology of the subgeneric taxa oi Anemone, Hoot et al. (1994) 
recognized two subgenera and seven sections. Section Anemone consists of 
four taxonomically unnamed groups, one of which (the Coronaria group) is 
characterized by distinctive tuberous rootstocks, typically heteromorphic 
1-3 ternate radical leaves, simple or branched scapes with sessile involucral 
bracts positioned near the middle of the peduncle, numerous linear to 
obovate sepals, and numerous, usually densely woolly achenes borne in 
ellipsoidal to cylindroidal heads. According to Hoot et al. (1994), the ca. 18 
species of the Coronaria group of section Anemone range from southern 
Europe to Mongolia, central and southern United States and adjacent 
Mexico, and South America below 10° south latitude (Hoot 1991; Joseph 
and Heimburger 1966). 

192 S]DA16(1) 1994 

Texas has five species of anemones with tuberous rootstocks: A . caroliniana 
Walter, A . berlandien Pritzel, A . tuberosa Rydberg, A . edwardsiana Tharp, and 
a new species, A. okennonii Keener & Button, herein described from the 
Edwards Plateau. The aim of our paper is to clarify the taxonomy of these 
species oi Anemone in Texas, to provide keys to the taxa, and to describe the 
new species. Joseph and Heimburger (1966) reviewed the cytotaxonomic 
structure of these species (except A. okennonii), and provided (p. 908) a 
helpful comparative chart and distribution maps; the reader is referred to 
their paper for additional details. The five species treated herein are 
distinguished chiefly on the basis of leaf dissection, scape pubescence, 
relative similarity of involucral bracts to the basal leaves, style length, and 
sepal number and size (see Table 1 for additional details). 

Until the species oi Anemone with tuberous rootstocks are studied world- 
wide, it would be premature to speculate on phylogenetic relationships of 
the species treated in our paper (See Hoot et al. [1994] for further analysis 
of this problem). We believe, however, that Anemone okennonii is taxonomi- 
cally closest to A. edwardsiana, but whether these two species are closest 
phylogenetically remains in question. 

. Involucre below middle of scape at antt 
long or longer than the ovoid achenes ... 

2. Scape usually branched, bearing 2 or more flowers per stem; involucral 
bracts similar or dissimilar to radical leaves; scape nearly glabrous to 
glabrous below involucre. 

3. Involucral bracts similar to radical leaves; radical leaflets narrow, 1-2 
cm wide; achenes 3-4 mm wide; rare in Texas (El Paso Co.) 


1. Anemone caroliniana Walter, Fl. Carol. 157-158. 1788. 

Slender, apically villous herbs; stems simple, 0.5-3.5 dm tall at anthesis, 
from rhizomes bearing a series of small, + oblong to globose tuberous 
rootstocks 0.5-1 cm long and 0.5 cm thick. Radical leaves 1-2 ternate, to 
3.5 cm wide; petioles 3-9 cm long. Leaflets 1-2 cm wide, glabrous, deeply 
2-3-parted or divided, segments variously cleft, lobed or toothed, ultimate 
segments broad to narrow with crenate-dentate margins to variously linear 
with acute tips, sessile to long petiolulate. Involucral bracts usually similar 
to at least some radical leaves, reduced, opposite or whorled, sessile to 
subsessile, sparsely pilose, positioned below middle of scape at anthesis. 
Flowers solitary; scape villous above involucre, nearly glabrous below. Sepals 
10-30, greenish white or pink to purplish blue, linear-oblong, 10-20 mm 
long and 1-6.5 mm wide; anthers yellow, 0.5-0.75 mm long. Fruiting 
heads ellipsoidal, to 2 cm long and 1 .5 cm broad; achenes brownish, turgid, 
±ovoid, ca. 2 mm wide, densely woolly; styles greenish (lavender-tipped), 
erect, filiform, as long as or longer than achenes, projecting beyond the 
woolly tomentum; 2n=l6 (Joseph and Heimburger 1966). 

February to April. Prairies, moist sandy soils, calcareous dry grasslands, 
rocky hillsides, and alluvial flats. South Dakota to Texas, east to South 
Carolina and Georgia. 

Anemone caroliniana is distinguished by its rhizomatous rootstocks bear- 
ing a series of small tubers, its heteromorphic radical leaves some of which 
resemble the involucral bracts, its simple scape bearing involucral bracts 
below the middle, its scape pubescent above the involucre but glabrous 
below, and its densely woolly ovoid achenes bearing relatively long styles 
(Table 1). In Texas, A. caroliniana occurs chiefly in the eastern third of the 
state, with some populations in the west-central and northern regions (see 
Joseph and Heimburger 1966, Fig. 3 , for a distribution map of this species). 

2. Anemone berlandieri Pritzel, Linnaea 15:628-629. 1841. 
A.hetenphylla^\xtt2.V\, nomen nudum insyn.,Torrey& A. Gray, Fl. N. Amer. 1:12. 1838. 
A. caroliniana Walter var. heterophylla Torrey & A. Gray, Fl. N. Amer. 1:12. 1838. 

A. decapetala Arduino var. heterophylla (Torrey & A. Gray) Britton & Rusby, Trans. New 

Similar to, though often larger than A. caroliniana. Stems simple, (1)3-5 
dm tall at anthesis, from thick clavate to oblong, tuberous rootstocks 2-4 
cm long and ca. 1 cm thick. Radical leaves 1(2) ternate; petioles 3-20 cm 
long. Leaflets 2-4 cm wide, pubescent, often 2-3 lobed or parted, the 
cuneate segments usually broad or rounded apically with dentate or serrate 
to crenate margins, sessile to long petiolulate. Involucral bracts dissimilar 
to radical leaves, ultimately dissected into linear segments with acute tips. 


pilose, positioned above middle of scape at anthesis. Flowers solitary; scape 
villous throughout. Sepals 7-17, greenish white to pink or purphsh blue, 
linear-oblong, to 7-1 5(20) mm long and 1.5-5 mm wide; anthers yellowish 
brown, 0.5-1 mm long. Fruiting heads cylindroidal, 2-3.5 cm long and 
0.5-1.3 cm broad; achenes flat, orbicular, 2-3 mm wide, densely woolly; 
styles greenish (lavender-tipped), subulate with bent tips, ca. 1/3 length of 
achenes, not projecting beyond the dense woolly tomentum; 2n^ 16 (Joseph 
and Heimburger 1966). 

February to April. Limestone hills, grassy knolls, and stony ground, Texas 
and Oklahoma eastward to (rarely) North and South Carolina. 

Anemone berlandieri is distinguished by its ternate radical leaves with 
broad segments having typically crenately toothed margins, its simple scape 
bearing involucral bracts above the middle of the more or less uniformly 
pubescent scape, its bracts distinctly unlike the radical leaves, and its 
densely woolly orbicular, flat achenes bearing styles ca. 1/3 length of the 
achene (Table 1). In Texas, A. berlandieri occurs throughout the state except 
for the extreme western portions (i.e., High Plains and Trans-Pecos, 
Mountain and Basin regions; see Map 1, Correll and Johnston 1970). 

The taxonomy of this species is complicated, owing largely to whether the 
North American plants are conspecific with the South American species 
recognized as A. decapetala Arduino (Britton 1891), and the adoption of the 
name A. heterophylla, which is listed by Torrey and A. Gray (Fl. N. Amer. 
1:12. 1838) merely as a synonym for A. caroliniana Walter var. heterophylla 
Torrey & A. Gray. Joseph and Heimburger (1966) clarified distinctions 
between the North and South American species, and Keener (1975) re- 
viewed the legitimacy of the name A. heterophylla. 

3. Anemone tuberosa Rydberg, Bull. Torrey Hot. Club 29:151-152. 

Robust, apically villous herbs; stems simple below, usually branched 
above mvokicre, 1-3 dm tall at anthesis, from brownish, oblong-obovate 
tuberous rootstocks 1.5-2 cm long and 1 cm thick. Radical leaves several, 
1-2 ternate; petioles 5-7 cm long. Leaflets 1-2 cm wide, subglabrous, 
variously parted or cleft, with cuneate-obovate segments having acute tips, 
sessile to petiolulate. Involucral bracts 3, similar to radical leaves in dis- 
section, reduced, short-petiolate, thinly pilose, positioned above middle of 
scape at anthesis; secondary branches 1 or more, 2-bracteate. Flowers 1 -several 
per scape; lateral flowering branches maturing later than central flower; 
scape villous above involucre, subglabrous below. Sepals 8-10, greenish white 
to purplish, linear-oblong, 10-14 mm long, 3-6.5 mm wide; anthers 
yellowish brown, ca. 1 mm long. Fruiting heads cylindroidal to ellipsoidal, 
1.5-3 cm long and 1-1.5 cm broad; achenes flat, orbicular, 3-4 mm wide, 
densely villous; styles lavender, ±erect, filiform, less than 1/2 length of 
achenes, not projecting beyond the dense woolly tomentum; 2«= l6 (Joseph 
and Heimburger 1966). 

April to May. High dry rocky slopes, southeastern California, southern 
Nevada, and southwestern Utah, southeastward to extreme western Texas 
(El Paso Co.), where it occurs on igneous rocky slopes (Joseph and Heimburger 

Anemone tuberosa is distinguished by its ternate to biternate radical leaves 
with acute-tipped oblong-ovate segments, its relatively large involucral 
bracts similar in dissection to the radical leaves, its typically branched scape 
with an involucre above the middle at anthesis, its scape pubescent above the 
involucre but more or less glabrous below, and its densely villous, flat, or- 
bicular, achenes having styles less than 1/2 the length of the achene (Table 1). 

Britton (1891) regarded the North American populations conspecific (as 
A. sphenophylla) with similar populations in Chile, but Rydberg (1902) 
pointed out that in the Chilean plants "the whole scape is decidedly 
pubescent with ascending hairs and the segments of the involucral bases are 
narrower." However, Lourteig (1951: 564) regarded A. sphenophylla as a 
taxonomic synonym of A. decapetala Arduino. 

4. Anemone edwardsiana Tharp, Amer. Midi. Naturalist 33:669- 1945. 
Slender, apically villous herbs; stems simple below, usually branched 
above involucre, 3-5 dm tall at anthesis, from brownish, oblong-obovate 
tuberous rootstocks 2-4 cm long and 1-2 cm thick. Radical leaves several 
to many, 1(2) ternate; petioles 8-15 cm long. Leaflets reniform in outline, 
2-2.5 cm wide, glabrous, variously 2-4 cleft, lobes crenate-dentate to 
sharply toothed, sessile to petiolulate. Involucral bracts 3, ±sessile, dissimi- 
lar to radical leaves, 2-5 cm long, ultimately dissected into oblanceolate 

segments with rounded to acute tips, glabrous to sparsely pilose, positioned 
above middle of scape at anthesis; secondary branches 1-3, 2-bracteate. 
Flowers 1 -several per scape; lateral flowering branches usually maturing 
with central flower; scapes villous above involucre, subglabrous below. Sepals 
10-20, greenish white to bluish, oblanceolate, 10-16 mm long, 1.5-3(4) 
mm wide; anthers bright yellow, 0.5-1 mm long. Fruiting heads ellipsoidal 
to cylindroidal, 1.5-3 cm long and 0.6-0.8 cm broad; achenes flat, orbicular, 
2-3(3.5) mm wide, typically sparsely pubescent to glabrous; styles erect to 
horizontal, filiform, ca. 1/3 length of achenes, not projecting beyond the 
woolly tomentum; 2;?= 16 (Joseph and Heimburger 1966). 

February to April. Alkaline soils in moist shaded canyons; shaded, moist, 
rocky limestone bluffs and ledges along eastern edge of Edwards Plateau 
(R.J. O'Kennon, pers. comm.). Two varieties: 

a. Achenes woolly, ±dull var eduarduana 

4a. An 

Range and habitat of the species. 
4b. Anemone edwardsiana var. petraeaCorrell, Madrono 19:189. 1968. 

Tall, spindly plants with glabrous, vernicose achenes endemic to moist 
rocky crevices along Curry Creek near Kendalia, Kendall Co., Texas. Addi- 
tional population and cytotaxonomic studies are desirable to establish more 
clearly the range of this taxon as well as its phylogenetic relationships to 
other species in this complex. 

The most polymorphic of Texas anemones, A. edwardsiana combines fea- 
tures of both A. tuherosa and A. berlandieri (cf. Table 1). It is like A. tuberosa 
in having a branched scape glabrous or slightly pubescent below the in- 
volucre, and like A . berlandieri in having involucral bracts with long, narrow, 
sharply toothed segments quite dissimilar from the radical leaves. Achene 
shape, style length, and position of the involucral bracts are similar to those 
character states in both A. berlandieri and A. tuberosa. The combination of 
branched scapes, involucral bracts dissimilar to the radical leaves, and 
relatively few sparsely woolly to glabrous achenes demarks A. edwardsiana 
from the other anemones in Texas. 

Joseph and Heimburger (1966) pointed out that A. edwardsiana not only 
combines features of A. tuberosa and A. berlandieri {= A. heterophylla in their 
treatment), but also "shows an intermediate geographic range and altitude 
preference." They suggested thdit A. edwardsiana may be a hybrid derivative 
of A. tuberosa and A. berlandieri. However, based on restriction site analyses, 
Hoot et al. (1994) claimed that A. edwardsiana is a sister species to A. 

berlandieri and possesses "no rDNA fra^^ments in common with A. tuberosa." 
Additional biosystematic studies are desirable to clarify the phylogenetic 
relationships of this complex. 

Because of glabrous, vernicose achenes, Correll (1968) segregated several 
populations occurring near Kendalia in Kendall Co., Texas, as A. edivardsi- 
ana vat. petraea Correll. These populations appear to be an ecotypic phase of 
the polymorphic A. edwardsiana. 

5. Anemone okennonii Keener & Dutton, sp. nov. (Figs. 1,2) 

Herba perennis erecta e tuberibus brunneis; tubera oblongo-obovata, 1-3 cm longa, 1 cm 
crassa. Folia radicalia pluria, triternata, glabriuscula; pecioli usque ad 10 cm longi. Foliola 

Slender, apically villous herbs; stems simple below, usually 1-3 branched 
above involucre, 2-3 dm tall at anthesis, from brownish oblong-obovate 
tuberous rootstocks 1-3 cm long and 1 cm thick. Radical leaves several to 
many, (2)5 ternate; petioles 5-10 cm long. Leaflets 0.3-1.5(2.3) cm wide, 
subglabrous but marginally ciliate, deeply 2-4 cleft, lobes cimeate-acnte, 
reddish-glandular; petiolules to 2 cm long. Primary involucral bracts 3, 
short-petiolate, dissimilar to radical leaves, 2-5 cm long, variously cleft into 
linear, acute-tipped segments, appressed pilose, positioned above middle of 
scape at anthesis; secondary branches (0)1-2(3), 2-bracteate. Flowers 2- 
several per scape; lateral flowering branches maturing later than central flower, 
scapes pubescent above involucre, glabrous or subglabrous below. Sepals 7- 
11, greenish white, oblong, 6-12 mm long, (2)3-4.3 mm wide; anthers pale yellow, 
fading to creamy brown, 0.75-1 mm long. Fruiting heads oblong-ellipsoidal, 
1-3 cm long and 0.5-1 cm broad; achenes flat, ovate, 2-3 mm broad, densely 
white villous; styles lavender-tinted, ±erect to horizontal, filiform, ca. 1/4 length 
of achenes, not projecting beyond the wooly tomentum. Chr( 
ber unknown. 

Representative specimens examined: U.S.A.: Texas: Brewster Co.: Glass Mts., 21 Mar 
l9A\,Rose-lnnes & Warnock 365 (SMU). Crockett Co.: 14.8 mi W of Ozona, l4 Mar 1949, 
Turner & Warnock 275 (SMU). Kimble Co.: Hwy 385, 2.5 mi S of Llano River, 9 Mar 1992, 
R.J. O'Kennon 8813 (PAC). Mitchell Co.: Lake Hollywood, 27 Mar 1945 , Pohl4744 (SMU). 
Pecos Co.: 20 mi NE of Ft. Stockton toward McCamey, 27 Apr 1 947, Warnock 5 1 99 (SMU). 


Kenner and DUTTON, Anemone okennonii 20 1 

Terrell Co.: 31 mi S of Sheffield, 14 Mar 1949, Turner & Warnock 507 (SMU); 4.2 mi W of 
Sanderson, 14 Mar 1949, Turner & Warnock 3 19 {SMU). Val Verde Co.: 20 mi NNW of Del 
Rio, 31 Mar 1947, McYaugh 7729 (SMU). 

March to April. Dry sunny rocky limestone ledges, hills, and roadsides, 
in red sandy loam and igneous-based soils at an elevation of 490-530 m. 
Endemic from Gillespie and Mitchell cos., southwestward to Val Verde and 
Brewster cos., Texas, mainly in the Edwards Plateau vegetational area with 
outliers in the Rolling Plains and Trans Pecos, Mountain and Basin vegeta- 
tional areas (see Map 1, Correll and Johnston 1970). 

Anemone okennonii has been variously recognized as A. caroliniana Walter, 
A. decapetala Arduino, A. edwardsiana Tharp, A. sphenophylla Poppig, A. 
tuberosa Rydberg, or possibly a hybrid between A . tuherosa and A . heterophylla 
Nuttall (= A. berlandien Pritzel). 

Anemone okennonii differs from its nearest presumed congener, A. edward- 
smna, by its leaf dissection ([2}3 ternate vs. 1|:2} ternate), its smaller leaflets 
(0.5-1.5 cm wide vs. 2-2.5 cm wide) with sharply cleft (rounded) lobes, its 
central flower maturing before (with) the lateral flowers, its sepal number 
(7-11 vs. 10-20), anther color (pale yellow vs. bright yellow), achene pu- 
bescence (densely white villous vs. sparse to glabrous), seedling leaves (similar 
to basal leaves vs. similar to bracts), relatively later blooming period (March 
to April vs. February to April), and habitat (drier, more exposed sites vs. more 
moist and shaded sites). The following chart summarizes differences be- 
tween A . okennonii and other Texas species oi AnemoneWixh tuberous rootstocks: 

10. Sepals 7-11 Uke A. tuberosa 

11. Sepals oblong, 6-12 mm X (2)3-4.5 mm Distinctive 
Plant associates of A. okennonii include Acacia roemeriana Scheele, Aristolo- 

chia coryi I.M. Johnston, Colubnna texensis (Torrey & A. Gray) A. Gray, 
Delphinium cawlinianum Walter, Draba cuneifolia Torrey & A. Gray, Echinocereus 
enneacanthus Engelmann, Euphorbia acuta Engelmann, Ferocactus setispinus 
(Engelmann) L. Benson, Gilia ngidula Bentham, Haplopappus spinulosus 
(Pursh) DC, Melampodium leucanthum Torrey & A. Gray, Opuntia phaeacantha 
Engelmann, Prunm minutiflora Engelmann, Tetraclea coulteri A. Gray, and 
Thamnosma texana (A. Gray) Torrey (R. J. O'Kennon, pers. comm.). 

Anemone okennonii appears to have evolved from A. edwardsiana and may 

actually be a xenc form of k (R. J. O'Kennon, pers. comm.). O'Kennon has 
suggested (pers. comm.) that millions of years ago when western Texas was 
more mesic than at present a precursor form of A. edwardsiana was most 
likely the predominant species o^ Anemone in this area. As western Texas 
became hotter and drier (and this trend continues at present), xeric forms 
began to evolve. Anemone tuberosa probably evolved first farther west, and 
then A. okennonii evolved to fit into the niche between the two. The three 
species seem not to overlap anywhere in their ranges. The only species 
sympatric with A. okennonii is A. berlandien, but there does not appear to be 
any intergradation between these two species. Critical biosystematic and 
population studies would be useful in establishing the most probable 
phylogenetic relationships among these five taxa of Texas anemones with 
tuberous rootstocks. 

Special thanks are due to Robert J. O'Kennon for pointmg out 
taxonomic problems in the Texas anemones and for providing specimens i 
data; to Dr. Sara Hoot and Dr. Ronald A. Pursell for reviewing an ear 
manuscript; and to Robert EC. Naczi for providing the Latin diagnosi: 
Anemone okennonii. Any errors of fact or interpretation are strictly ^ 

Britton, N.L. 1 89 1 . The American species of the genus Anemone ar 

id the ^ 

been referred to it. Ann. New York Acad. Sc. 6:215-238. 

CoRRELL, D.S. 1968. Some additions to the flora of Texas— IV. Mad, 

rono IS 

): 187-192. [Type 

description oi Anemone edwardsiana Tharp var. petraea Correll, 1 


r plant 

:s of Texas. Texas 

Research Foundation, Rennet. 
Hoot, S.B. 1991. Phylogenetic relationships in Anemone based on morphology and 

chloroplast DNA variation. Ph.D. dissertation, Univ. of Michigan. 
, A.A. Reznicek and J. Palmer. 1994. Phylogenetic relationships in Anemone 

(Ranunculaceae) based on morphology and chloroplast DNA. Syst. Hot. 19:169-200. 
Joseph, C. and M. Heimburger. 1966. Cytotaxonomic studies on New World species of 

Anemone (Section Eriocephalm) with tuberous rootstocks. Canad. J. Bot. 44:899-928. 
Keener, C.S. 1975. Studies in the Ranunculaceae of the southeastern United States. I. 

Anemone L. Castanea 40:36-44. 
LouRTEiG, A. 1951. Ranunculaceas de Sudamerica templada. Darwiniana 9:397-608. 
Rydberg, P.A. 1902. Studies on the Rocky Mountain flora— VII. Bull. Torrey Bot. Club 

29:145-160. {Type description oi Anemone tuberosa Rydberg, 151-152.} 



Department of Botany 

University of Texas 

Austin, TX 78713 U.S.A. 

Jaimehintonia is a recently described monotypic genus of the Amarylli- 
daceae, tribe Allieae (Turner 1 993). The single species,/ gypsophila, is a rela- 
tively rare edaphic endemic in to gypseous soils of northeastern Mexico. Its 
generic relationships are moot, but its author thought the species to be most 
closely related to Androstephium and allies, in the sense of Moore (1953). 

The present paper attempts to position the genus with more certainty 
using karyotypic data. 

ies were made from root tips of seeds germinated on a 
agar nutrient medium in a petri dish. Seeds were collected at the type localit 
by George S. Hinton (Wnton et al. 20560, TEX). 

Root tips were placed in saturated 8-hydroxyquinoline solution for 4 h 
fixed in 3 : 1 (ethanol : acetic acid) about 4 hr, and transferred to 70% ethane 
overnight. They were softened in 10% HCl at 60 C for 15 min, the 
squashed using standard procedures. 

The chromosome number of this species is 2n = 20 (Fig. 1). Its karyotype 
can be described as follows (Fig. 2): three pairs of large submetacentric 
chromosomes, three pairs of medium metacentric chromosomes, three pairs 
of small metacentri 

Jaimehintonici appears to have a distinctive karyotype that, along with its 
morphological features, supports its recognition as a distinct genus. Among 
the tribe Allieae, chromosome numbers of 2n = 20 are found only in two 
species of the genus Muilla, but chromosomes of the latter differ in shape 
from those oi Jamiehintonia. (Both M. maritima [Torr.} S. Wats, and M. 
trammontana Greene have five medium to long submetacentric chromo- 
somes, four short submetacentric, and one short nearly metacentric chromo- 
somes; Lenz 1975.) An anomalous count of n = 20 pairs has been reported for 
Brodiaea lutea (Lindl.) Morton var. anilina (Greene) Munz (Niehaus 1965), 
but the chromosome numbers of most taxa oi Brodiaea are based on x = 6, 7, 
8, and 9 (Federov 1969). This, along with differences in floral morphology, 
appears to rule out a close relationship hev^een J aimehintonia and Brodiaea. 
In contrast, the similarity between Jaimehintonia and taxa of the An- 
drostephium group (within the Allieae), as noted by Turner (1993), indicate 

Zhao, Karotype of J. 

*,- ft 

'1 u n 

%% ll S It fi § 

nary relationship is probable. Unfortunately, i 
nber reports for Androstephium. 

1 grateful to Dr. Guy Nesom and Dr. B.L. Turner for helpful sugges- 
1 preparing the manuscript, and to Dr. Richard Starr for providing the 
It medium. 

Fedorov, a. a. (ed.). 1969- Chromosome numbers of flowering plants. Acad. Sci. U.S. S.I 

iotMutlla coronata and M. transmontana. Aliso 8:259-262. 
Mcx)RE, H.E., Jr. 1953. The genus Muilla and its allies. Genres Herb. 8:263-294. 
NlEHAUS, T.F. 1965. In: Documenred chromosome numbers of plants. Madroiio 1 



Belousova, L.S. and L.V. Denisova. 1992. Rare Plants of the World. 
(ISBN 90-6191-482-5, hbk). A.A. Balkema Publishers, Rotterdam, 
Netherlands. $75 (hbk). 348 pp. 

plant than is available from the Red Data Book of Plants published by the World 
Conservation Union (lUCN). Selections are based on regional publications as well as the Red 

North America). When examining endangered, threatened, or protected native plants of 

Reichenbach's echinocereus {Echinocereus rekhenbachti) and Tobusch's ancistrocactus 
{Anastrocactus tobuschii). Hinckley's oak {Quercus hinckkyt) and Texas Wild-Rice {Ztzama 


our attention that the name Karinia Reznicek & McVaugh, Flora Novo- 
Galiciana 13:386. 8 Dec 1993, may not be validly published because, even 
though monotypic, the sole species was not stated in the protologue also to 
be the holotype of the generic name. 

It is difficult to understand how the generic name could have any other 
type, but for the peace of mind of those who would consider the name not 
validly published, the name is here explicitly typified to fulfil this require- 
ment for valid publication, and the combination Karinia mexicana is remade 
now that the generic name is valid. 

Karinia Reznicek & McVaugh 

— A. A. Reznicek & R. McVaugh, University of Michigan Herbarium, North 
Umversity Building, Ann Arbor, Ml 48109, U.S.A. 

TEXAS — Penstemon thurberi Torr. (P. scoparius A. Nels., Leiostemon thurberi 
Greene, P. ambiguus var. thurberi Gray) is a woody perennial distributed from 
Southern California through Arizona to Southcentral New Mexico and 
Northern Mexico. In New Mexico, it is found in the southwestern counties 
east to Dona Ana and Lincoln counties (Martin and Hutchins 1981). This 
is the first report of P. thurberi in Texas, a range extension of approximately 
320 km from the nearest known site in Dona Ana county, New Mexico, to 
the two populations on the slopes of Little Pinto Canyon and Cinco de Mayo 
Canyon in the Chinati Mountains of Presidio county, Texas. The two 
populations, approximately 2 km apart, one with about 150 specimens and 
the other with about 300, are in loose sandy shale, surrounded by plants 
typical of the Chihuahuan Desert grassland. 

Collection data are: TEXAS. Presidio Co.: Little Pinto Canyon, "^armck 121, 9 Aug 
1988 (Fig. 1). 

— Barton H. Warnock, Professor Emeritus, Department of Biology, Sul Ross State 
Umversity, P.O. Box 158, Alpine, TX 79851, U.S.A.; John Mac Carpenter, 
Native Plant Society of Texas, P.O. Drawer 430, Ft. Stockton, TX 79735-0430, 

Martin, W.C. 

1982. A flora of New I 

MDLEY. 1915. Flora of New ^ 
rinting Office, Washington. 

FLORIDA — Euphorbia graminea)sicq. apparently c^me into southemFlonda. 
during the late 1980s as a weed in horticultural stock. John Popenoe found 
plants without flowers in nurseries for several years (personal communica- 
tion), but it was not until Jan-Feb 1993 that Popenoe and I independently 
found flowering populations ofE.grammea separated by several miles. Plants 
of this species were found again, and in new areas, during the fall of 1993 
so it seems fair to regard the species as established in southern Florida. 

Euphorbia graminea is an erect annual reaching 35 cm in height with leaves 
well separated along the stem and branches. Stem leaves are 2-4 cm long, 
alternate and, on all Florida plants seen, are ovate with undulate margins. 
Leaves on the branches of the inflorescence are slightly shorter, opposite or 
nearly so and narrowly elliptic. Cyathia are terminal or solitary in the axes 

of dichasia and about 1 mm long. A variable number (1-4) of glands is pro- 
duced along the upper edges of the cyathium, with many specimens having 
two glands on the cyathium. Glands have white appendages that are about 
equal in size to the glands. The capsule is held beneath the gland appendages 
until maturity and may not be apparent without close inspection. Mature 
capsules are about 3 mm long and wider than long. 

Euphorbia graminea is widespread from southern Mexico to northern South 
America, and has weedy tendencies throughout this range. It shows consid- 
erable variation in leaf shape and pubescence, leading to the proposal of 
several subspecific groups. There is no consensus on the validity of these 
subspecific taxa (Webster and Burch 1968), so no attempt was made to 
identify the Florida plants below the specific level. 

Within the genus Euphorbia (sens, lat.), E. graminea is the lectotype species 
of the section Cyttarospermum (Wheeler 1943). It has also been treated under 
the segregate genus Eumecanthus. This group is characterized by petiolate 
leaves and ecarunculate seeds with an areolate pattern of raised bumps 
covering the surface (Subils 1977). 

All collections from Florida so far have been made in cultivated sites. 
Within these cultivated areas, the Euphorbia evidently prefers bare patches. 
Seeds are dispersed locally by the explosively dehiscent capsules, but the pri- 
mary means of spread in southern Florida seems to be through human trans- 
port of ornamental plants, especially field grown trees. 

Voucher specimens: FLORIDA. Dade Co.: weed in residential lawn, near Homestead, 28 
Jan 1993, Herndon 3338 (FTG); weed at the Dave Romney farm, Homestead, 5 Feb 1993, 
Popenoe 2476 (FTG); growing in planted bed along the right-or-way of Old Cutler Road, ca. 
1/4 mi S of Fairchild Tropical Garden, 20 Feb 1993, Popenoe 2480 (FTG); weed in potted 
plant by residence, Perrine, 20 Aug 1993, Herndon 3346 (FTG). 

I am grateful to John Popenoe for sharing his knowledge of the early 
history of this species in Florida. — Alan Herndon, Department of Biological 
Sciences, Florida International University, Miami, EL 33199, U.S.A. 

Subils, R. 1977. Las especies de Euphorbia de la Republic 
Webster, G.L. and D. Burch 1968. Family 97. Euphorb 

Panama, Part VI. Ann. Missouri Bot. Gard. 54:211- 
Wheeler, L.C. 1943. The genera of the living Eupho 


210 SiDAl6(l) 1994 

FOR COLORADO— Me/zc^ L., a genus of ± 80 species, is found in tem- 
perate regions throughout the world, except Australia (Clayton and Ren- 
voize 1986). Species of Melica are found in a variety of habitats, from 
woodland shade to dry stony slopes (Clayton and Renvoize 1986). There are 
17 species of Meiu^a in the United States. Boyle (1945) reported three taxa 
as occurring in Colorado, M. bulbosa Geyer ex Porter & J. M. Coulter, M. 
porteri Lamson-Scribner var. porteri, and M. spectabilts Lamson-Scribner. In 
the summer of 1992, Stanley and Gretchen Jones discovered a fourth taxon 
in the state, M. suhulata (Grisebach) Lamson-Scribner. This represents a 
range extension of approximately 400 km (250 miles) from the closest 
previously known locality in Fremont County, Wyoming, as reported by 
Hallstenetal. (1987). 

Melka subulata has not previously been reported as occurring in Colorado 
(Boyle 1945; Chase 195 l;Harrmgton 1964; Weber 1987, 1990; Weber and 
Wittmann 1992). This species is found in mesic forests from Alaska south 
to Canada, Washington, Oregon, California, Idaho, Montana, Wyoming, 
and Colorado. 

The following is a key to identify the species of Melka found in Colorado. 

eins; palea about 2/3-3/4 as long 

3. Second glumes 2/3-3/4 the length of the spikelet M. bulhosa 

Voucher specimen. COLORADO. Gunnison Co.: Gunnison National Forest, 0.6 mi S 
on USFR 898 from McClure Summit, hillside seep in aspen-spruce forest with gray silty clay 
loam soil, elev. 2660 m (8727 ft), 24 Jul 1992, S. & G.Jones 9509 (jkw, sdj, TEX, UTC). 
Associates: Veratrum californkum Durand, Zigadenus elegans Pursh, Carex stenoptila F. Herm., 
C. microptera Mack. , Senecio bigelovii A. Gray, Rudbeckia occidentalis Nutt. , Mertensia sp. , Poten- 
tilla spp., Rosa sp., Delphinium sp., Geranium sp., Populus tremuloides Michx., and Picea sp. 
We would like to thank Mary E. Barkworth (UTC) for her review of this 
manuscript and Gretchen D. Jones (USD A) for her assistance with the field 
work.— Joseph K. Wipff and Stanley D.Jones, S.M. Tracy Herbarium, Department 
of Rangeland Ecology and Management, Texas A&M University, College Station, 
TX 77843-2126, U.S.A. 

BoYi.E, W.S. 1945. A cytotaxonomic study of the North American species of Melia 

Madrono 8:1-32. 
Crash, A. 1951. A revision of A. S. Hitchcock's manual of the grasses of the United State: 

2nd ed., U.S.D.A. Misc. Pubi. No. 200. U.S. Government Printing Office, Washing 

ton D.C. 
Clayton, W.D. and S.A. Renvoize. 1986. Genera graminum, Grasses of the World. Ke^ 

Bulletin Additional Series XIII. Her Majesty's Stationary Office, London. 
Hallsten, G.R, Q.D. Skinner and A.A. Beetle. 1987. Grasses of Wyoming. 3rd Editior 

University of Wyommg Research Journal 202. Agricultural Experiment Station, Larami< 

Harrington, H.D. 1964. Manual of the plants of Colorado. 2nd ed. The Swallow Press, In( 

Chicago, Illinois. 
Weber, W.A. 1987. Colorado flora: Western Slope. Colorado Assoc. Univ. Press, Niwoi 

Weber, W.A. 1990. Colorado flora: Eastern Slope. Univ. Press of Colorado, Niwoi 

a small tropical and subtropical genus of ten species, that is principally 
found in the New World (Clayton and Renvoize 1986). Three species are 
now known to occur in Mississippi: Eustachys caribaea (Sprengel) Herter, E. 
glauca Chapm. and E. petraea (Sw.) Desv. Eustachys caribaea, native to South 
America, has been reported from Louisiana (McKenzie et al. 1 987) and Texas 
(Wipff and Hatch 1 992). Eustachys glauca, native to North America, has been 
reported from Florida, Georgia and North Carolina and is found in brackish 
marshes, wet prairies and swamps (Chase 195 1). Eustachys petraea is found in 
the SE U.S.A., eastern Mexico, Belize, Honduras, Costa Rica, and Panama 
(Pohl 1 980). In the United States, E. petraea occurs in the Coastal Plain from 
North Carolina west to Texas on coastal sands, sandy fields and open pine 
woods (Chase 1951). 

fertile) floret conspicuously pubescent on the midvein and margir 
glume apex is conspicuously bifid, the lobes are rounded to acui 
ertile) lemma dark chocolate brown to black or pale to golden bi 

2(1). Lower (fertile) lemma dark chocolate brown to black; lower 1 


margms glabrous on the lower 1/2-2/3; the upper portion of the 1 
margins with short usually appressed trichomes 0.1-0.4 mm long; 
most (sterile) lemma 0.8-1.0 (1.2) mm long 


2. Lower (fertile) lemma pale to golden brown; lower lemma m 


pubescent the entire length; trichomes on the lower lemma m 
lemma 1.3-1.5 mm long 


Eustachys caribaea (Sprengel) Herter 

Voucher specimens. MISSISSIPPI: Harrison Co.: Gulfport, W jet. of Hwy 49 and 19th 
Street, T8S, RllW, S4; along edge of road and in a vacant lot, 11 Aug 1993, C.T. Bryson 
12661 (SWSL); Gulfport, W of Hwy 49, between 34th and Madison Street; vacant lot, open 
area, on sandy soil, 12 Aug 1993, C.T. Bryson 12662 (SWSL). 

Eustachys glauca Chapm. 

Voucher specimens. MISSISSIPPI. Hancock Co.: 1 mi S jet. Hwys US 90 and MS 607; 
NW of US 90; open roadside on humic sandy soil, 17 Oct 1993, C.T. Bryson 13181 & R. 
C^f^er(IBE,SWSL,TAES).HamsonCo.:Gulfport,MillRd.;T7S,RllW,Sec.25;old field 
adjacent to Bayou Bertrand, 1 1 Oct 1990; TC. Lockley s.n. (SWSL). 
—].K. Wipff, Stanley D.Jones, SM. Tracy Herbarium, Department of Rangeland 
Ecology and Management, Texas A&M University, College Station, TX 77843- 
2126, U.S.A. and Charles T Bryson, USDA-ARS, Southern Weed Science 
Laboratory, Stoneville, MS 38776, U.S.A. 

Chase, A. 1951. Hitchcock's 


Misc. Publ. No. 200. U. ; 

5. Gov 

Clayton, WD. and S.A. Rn 


Bull. Addit. Ser. XIII. He 

.r Maj, 



POHL, R.W 1980. Gramineae. In: Flora Co: 

Fieldiana,Bot. 4:1-608. 
Wipff, J. K. and S.L. Hatch. 1992. Eustachys , 

TUCKY— Field searches for rare native plants on the Daniel Boone Na- 
tional Forest in east central Kentucky are conducted yearly. During such a 
search in fall 1991, an unusual vine was found in McCreary County. The 
vine, growing with Parthenocissus quinquefolia and at first mistaken for an 
aberrant form of the latter, was eventually identified as Schisandra glabra 
(Brickell) Rehd., starvine. The plant was growing on a shale/sandstone talus 
slope at a break in a sandstone cliff. The species is typically a high climbing 
vine up to 3 cm in stem diameter, but the plants in this population were all 
creeping along the ground, some rooting at nodes. The site was partially 
shaded but did receive direct east sun. Overstory in the area consisted of Acer 
rubrum, Liriodendron tulipifera, Tsuga canadensis, and scattered Quercus species. 
Other species immediately adjacent to the site included Clethra acuminata, 
Kalmia latifolia, Mitchella repens, and Solidago caesia. 

Schisandra glabra was not reported for McCreary County by Rogers (1941) 
or for Kentucky by Braun (1943). More recently, the taxon was not reported 
for Kentucky by Johnson and Nicely (1990) or Browne and Athey (1992). 
Medley (1993) referenced my collection in his dissertation. This new 
location is about 250 km disjunct from the nearest population of 5". glabra, 
in Stephens County, Georgia Qones and Coile 1988; Ettman 1980) and 
represents the first record for the northern Cumberland Plateau. Other sites 
are from the southern Cumberland Plateau (Alabama), the Piedmont 
Plateau, the Coastal Plain, and the Mississippi Embayment. 

The leaves of starvine are alternate, typically pale green, somewhat fleshy, 
and coarsely and remotely serrate. Shorter branch stems have leaves crowded 
at the branch tip, almost appearing as palmately arranged leaflets. Leaves on 
trailing stems are widely spaced. Flowers are difficult to see as they are 
usually high above the ground. Fruits are scarlet and are arranged in loose 
racemes on pendulous peduncles. Illustrations may be found in Duncan 
(1967) and Stone (1968). 

I thank Michael A. Vincent (MU) for providing helpful comments and 

Voucher specimen: KENTUCKY. McCreary Co.: single population on shale/sandstone 
talus slope at break in sandstone cliff on W side of Wolfpen Branch, ca. 1 mi S of White Oak 
Junction, ca. 1 mi E of Hickory Knob Church, Barthell Quadrangle, elev. ca. 1220 ft, 8 Oct 
1991, D.D. &. LA. Taylor 16331 (BEREA, KNK, MU, US). 

—DavtdD. Taylor, USD A Foresi Service, Darnel Boone National Forest, 1833 
BtghillRoad, Berea, KY 40403, U.S.A. 


published by the author, Cincinnati. 
Browne, E.T. and R. Athey. 1992. Vascular plants of Kentucky: i 

The University Press of Kentucky, Lexington. 
Duncan, W.H. 1967. Woody vines of the southeastern United Sta 
Ettman, D. 1980. A study oiSchtsandra glabra Brickell) Rehder, a : 

Johnson, G.P. and K.A. NrcELY. 1990. The Magnoliales of Kenti 

Jones, S.B. ai 

nd N.C. CoiLE. 

1988. The distrib, 


t of Botany, Uni^ 

.ersity of Georgia,. 

Medley, M.E. 


ted catalog of knoNv 


^d dissertation, L 



1941. The flora of 

■McCreary County, 


of Kentucky, Lex 


Stone, D.E. 

1968. Cytologic. 

il and morphologic 

r flora of Georgia, 
r flora of Kentucky. 

tions in northern Cahuila, Mexico, have included two species not previously 
reported for the state. Nesom (1992) found a similar situation for three 
species of Aster collected in the Serranfas del Burro in Coahuila. 

Thlaspi montanum L. var. motitanum grows on moist or dry, open, 
rocky scree or talus slopes, alluvial fans or flats, limestone cliffs, and forest 
clearings. It has a wide distribution in the western United States, where it 
occurs from Washington, Oregon and southward along the Rocky Mountain 
Cordillera into New Mexico, Arizona, western-most Texas and in one locality 
in the La Bufa Mountains of northern Chihuahua, Mexico. This new record 
extends its distribution to the northwestern Coahuila. 

In addition to its wide geographical distribution, this variety of Thlaspi 
montanum has wide morphological variation (Holmgren 1971). The plants 
growing in Coahuila are 20-30 cm tall with racemes 8-10 cm long, peduncles 
1 cm long and fruit 1 cm long and 5-6 mm wide. The habitat is semi-mesic 

Specimens examined: MEXICO: Coahuila: Mpio. Muzquiz: Sierra del Carmen, Ranchos 

1300 m, 27 Mar 1992, AT. A. Carranza 1383, J. Noriega, y L. 
Mpio. Muzquiz: Sierra la Encantada, cucsta Malcna, aporx. 170 V 

Fraxmus cmpidata, Ungnadia speciosa, Quenus graves u y Q. mvaginata, 1 6()() m, 29 Mar 1 992, 
Af. A. Carranza 1525, J. Noriega y L. Garcia (ANSM). 

Oenothera missouriensis Sims. 

Specimens examined: MEXICO: Coahuila: Mpio. Muzquiz: Carr. Muzquiz-Boquillas 
(53), Rancho la Babia, area cercana ai arroyo la Babia, 28°33'N, 102°05'W, matorral de 
Lmcophyllum Futescens, Acacia rigidtda con Rhm virens, R. lanceolata y Ephedra, 950 m, 17 Sep 
1992 J. A. Willarreal 6971, M. A. Carranza y R. Vasquez A. (ANSM, MEXU). 

Oenothera missouriensis is widely distributed in limestone knobs, prairies 
and dry hills in the High Plains through Oklahoma, west Arkansas, 
Missouri to Kansas and Nebraska and into northcentral Texas (the Edwards 
Plateau). The coahuilan population belongs to the var. incana gray and is 
located in an intermountain valley in central Coahuila disjunct horn the 
closest Texan location. Numerous other species show a similar pattern of 
distribution as noted by Nesom (1992), considering that the two species 
here recorded have a primarily North American distribution extending into 

I thank G. Nesom for comments on the manuscript.— >jM. ViUarrealQ., 
Universidad Autonoma Agraria Antonio Narro, Departamento de Botdnica, 
Buenavista, Saltillo, Coahuila 25313, Mexico. 

:oRRELL, D.S. and M.C. Johnsi 
Research Foundation, Rennet 

loLMGREN, P.K. 197 1 . A biosyst 
Its allies. Mem. New York Be 

Jesom, G.L. 1993. Three spec 
Coahuila, Mexico. Phytologia 

IN ARKANSAS— Q/^m/j grayoides Mohlenbrock is an obscure member of 
Cyprus section Laxiglumt. It has previously been reported from Illinois, 
Missouri, Texas, and Louisiana. In Illinois its habitat is dry sand prairie 
located on the terraces of outwash plains of the Mississippi and the Illinois 
rivers (White & Madany 1978). In Missouri it is locally abun-dant on the 
Scotco sandndges of the southeastern lowlands (Carter & Bryson 1991). The 
Texas and Louisiana occurrences are described as located on pine barrens, 
xeric riparian sandhills, sandhill woodland, and deep sand savanna (Bridges 
& Orzell 1989). An Arkansas station was discovered as follows: 

NWl/4ofNEl/4Section6, T12N, R19W, onAgalasoil, 11 Sep \99i, Logan 1993-93 

Approximately 100 plants were observed in a sunny opening of a pine 
sandhill community. Associate species mciude Polygoneila americana, Stylisma 
pickeringii, Quercus incana, Q. stellata var. margaretta, Polansia erosa, Hapiopa- 
ppus divaricatus, and Monarda punctata. The site is undergoing active erosion. 
The presence of active disturbance to the site corresponds to descriptions 
given for sites in other states (Bridges & Orzell 1 989; Carter & Bryson 1 99 1 ; 
Herkert 1991)- The pine sandhill community where the species was found 
corresponds more closely to the descriptions oiC.grayoides locations in Texas 
and Louisiana (Bridges & Orzell 1989) than to those in Missouri or Illinois 
(Carter & Bryson 1991). 

Discovery of C grayoides in Arkansas indicates that this species should be 
looked for in other sandhill communities of southwestern Arkansas. Addi- 
tionally, the presence in northeastern Arkansas of soils derived from sand de- 
posits related to the sandy soils of southeastern Missouri (Saucier 1978) in- 
dicates that the species should also be searched for in that portion of the state. 

I am grateful to the following individuals for their assistance: Dr. Richard 
Carter, Valdosta State College; Dr. Paul McKenzie, U.S. Fish and Wildlife 
Service; and Sherry Holmes, Missouri Department of Conservation. — -John 
M. Logan, Arkansas Natural Heritage Commission, 1500 Tower Building, 323 
Center St., Little Rock, AR 72201, U.S.A. 

IRIDGES, E.L. and S.L. Orzell. 1989. Additions and noteworthy vascular plant collections 

from Texas and Louisiana, with historical ecological and geographical notes. Phytologia 

;arter, R. and C.T. Bryson. 1991. A report oi Cyperus grayotdes and Cyperus retroflexm 

(Cyperaceae) new to Missouri and notes on other selected Missouri Cyperus. Sida l4: 

Ierkert, J.R. (Ed.) 1991. Endangered and threatened species of Illinois: status and 

distribution Volume 1: plants. Illinois Endangered Species Protection Board. 
AUCIER, R.T. 1978. Sand dunes and related eolian features of the lower Mississippi River 

Alluvial Valley. Geosci. and Man 19:23-40. 
^HiTE, J. and M. Madany. 1978. Classification of natural communities m Illinois. In: 

White, J. ed. Illinois Natural Areas Inventory Technical Report. Vol. I: Survey Methods 

and Results, p. 311-405. Illinois Department of Conservation. 


Horn, B., R. Kay and D. Abel. A Guide to Kansas Mushrooms. (ISBN 
0-7006-0570-3, hbk; ISBN 0-7006-0571-1, pbk.). University Press 
of Kansas, 2501 West 15 th Street, Lawrence, KS 66049-3904. (91 3)864- 
4154. $19.95 (pbk); $29.95 (hbk) 298 pp., 6 figures, 160 color 

Kansas is famous for many things; wheat, cattle, oil, gas, aircraft manufacture, politi- 
cians, basketball, mile runners and now you can add mushrooms to the list. Most people not 
native to Kansas who plan a trip using Interstate 70 conjure up images of a monotonous, 
flat landscape dominated by endless fields of wheat. Nothing could be further from the truth 
for eastern and central Kansas. The westernmost extension of the broadleaf deciduous forest 
reaches into eastern Kansas, providing diverse habitats for mushrooms. The Flint Hills in 

heavily wooded with a mixture of deciduous trees and junipers. These diverse habitats 
provide for a much greater biodiversity of flowering plants, mushrooms, Myxomycetes, 

Bruce Horn, Richard Kay, and Dean Abel have given us a book that documents over 750 

bstrates, sex and mushrooms, mycology in Kansas, first fungi, 
ndar of fungal fruitings, collecting mushrooms, photograph- 

ation is available in other mushroom book 

s but some is not. The section on 

s destroys the myth that Kansas is a flat desi 

ert. First fungi informs the reader 

' to recognize groups of fungi. A Kansas 

! calendar of fruitings gives the 

:currence and the months collected for 15 

fungal species. There is a very 

1 collecting mushrooms. There is 

36 named Gigantomyces kamensts, depicted 

in color and located in Mushroom 

• Kanopolis Lake. A Kansas historical mark 

:er along Interstate 70 reveals the 

rue origin of these "mushroom rocks. "Formed through millions of years of erosion, and 
nown geographically as rhe Smoky Hills, these Dakota sandstone mounds cover a wide 
rea. Notice the natural rock outcroppings and toadstools" in this park. Here also are srone 
ence posts still used in this portion of Kansas." 
There is an interesting chapter on the history of mycology in Kansas that highlights the 

ournal of Mycology, the forerunner oiMycologia, that today represents the official publication 
f the Mycological Society of America. Elam Bartholomew who was a farmer near Stockton 
n Rooks County and a self taught mycologist, "...became the greatest collector of American 
angi in his generation." The numbers of fungi he collected are staggering: "...292,000 

218 SiDA 16(1) 1994 

Clark T. Rogerson and Robert L. Shaffer, currently distinguished mycologists, collected 
thousands of fungal specimens for the Kansas State University Herbarium and began an 
annotated checklist of Kansas fungi. But there is much more packed into these six pages. 
An Anthology of Kansas Mushrooms is divided into the Basidiomycotinaand Ascomycotina 
with keys to orders, to families, and to species. The dichotomous keys are based on 
macroscopic features and are easy to use. The authors are to be commended for giving 
numbers along with species names in the keys that guide the user to the species descriptions 
and photographs. Species are presented in systematic order and with the non technical 
species description and color habit photograph on the same page The color habit 

This book is well organized and easy to use for the beginner but has sufficient detail to 
tisfy the advanced student. There is a wealth of informarion not found in many regional 
ushroom books that warrants buying this book. It is the best of the regional books on 

Dick-Peddie, William A. 1993. New Mexico Vegetation, Past, Present, 
Future. (ISBN 0-8263-1 361-2, hbk) University of New Mexico Press, 
Albuquerque. $ 244 pp. + map and color photographs. 

New Mexico has a richly diverse vegetation that is in a continual state efflux due to both 
human influences and long-term climatic changes. This theme pervades New Mexico 
Vegetation in which Dr. Dick-Peddie summarizes and synthesizes data from his long career 
at the University of New Mexico. Not only does this volume provide a detailed account of 
the present vegetation of New Mexico, but also gives an overview of the physical factors, 
vegetational history, and future concerns about rare species and fragmented natural areas. 

The book is divided into twelve chapters which fall into three natural sets. Chapters One 
to Four, plus the Introduction, provide the reader with a general background on the 
vegetation. In particular, immediately preceding Chapter One there are a county map and 
29 handsome, colored plates that depict the vegetation types and phenomena discussed. 
Chapter One, covering the physical environment, includes maps of the physiographic 

ration, and soils. Chapter Two provides a rec 

the following vegetation types in New Mexico: Alpine Tundra, Subalpine Coniferous 
Forest, Upper Montane Coniferous (mixed conifer) Forest, Lower Montane Coniferous 
Forest, Aspen Disturbance Forest, Coniferous Woodland, Mixed Woodland, Savanna 
(exrensive woodland-grassland ecotone), Montane Grassland , Plains-Mesa Grassland, Desert 
Grassland, Montane Scrub, Plains-Mesa Scrub, Great Basin Desert Scrub, Chihuahuan 
Desert Scrub, Successional Montane Scrub, and several riparian vegetation types. Chapter 
Ten discusses special types including aquatic, lava-flow, and gypsophilic vegetation. W.H. 

The final chapters shift the focus from the description of presenr vegetation types to 
concerns of and factors effecting the future ot New Mexico vegetation Richard Spellenberg 

of limited distribution, and species in decline Besides reviewing the problems, needs, and 

species involved. In Chapter 1 2, Dick-Peddie ad\ .inces a \ ica\ of the future for the vegetation 

as a whole. In particular, his advocates a series ol natural aicas ind a means to secure them. 

When I first saw this book, I knew that it would W invaluable as a lecture and student 

somewhat disappointed when the book was c\ ikiated against the rigorous demands that I 
placed upon it. I was happily composmg charts ol \etic ration on a grid of mean annual 

ography of the Southwest. — Roger \1 

, Botanical Research Instit. 

Neubert, H., W. Nowotny and K. Baumann. 1993. Die Myxomyceten 
Deutschlands und des angtenzenden Alpenraumes unter 
besondererBemchschtigungOsterreichs. Band l,Ceratiomyxales, 
Echinosteliales, Liceales, Trichiales. (ISBN 3-929822-00-8, hbk) 
Karlheinz Baumann Verlag, Gomaringen, Germany. DM190 (ca. 
$110.00). 344 pp., 158 line drawings, 96 SEM, 192 colored photo- 
English translation of the title, The Myxomycetes of Germany and its bordering Alpine 

Areas, with special attention to Austria. Volume 1 Ceratiomyxales, Echinosteliales, 

Liceales, Trichiales. 

volumes. Volume I is 344 pages, bound in cloth with a color jacket, 190 German marks and 
about $110 at current exchange rates. 

Introductory pages have photographs of colored plates taken from "BuUiard's Histoire 
des Champignons de la France" that illustrate 5'/^OT0«/ ■ 

Trkhia contorta, Trichia decipiens and Arcyria incarnata. 
way the classification of the Myxomycetes, give their occurrence and distribution, describe 
and illustrate their life cycle, and discuss the general morphology of the fruiting bodies. The 
color photographs of the fruiting body habits are the highest quality I have ever seen. The 
book is worth the cost just for the excellent color photographs and scanning electron 
micrographs. Some of the color photographs are of fruiting bodies in an immature and 
freshly matured state. This is true oi Ceratiomyxa, Dktydiaethalium plumheum, Hemitrkhia 
davata, Lycogala epidendrum, Trkhia decipkm var. olivacea, and Tubifera ferruginosa. The 
illustrations of color photographs and line drawings are conveniently included in the text 
on the same page as the species description. The SEM's are included as a group at the back 

number cited as the source of the illustration. The authors are to be commended for 

A special feature of this book is the effective use of boldface headings that guide the reader 
through the genera, species and species descriptions. The high quality paper, double column 
format and accessibility of illustrations on the same page as the species description make this 
book very user friendly. 

All three volumes are in German and represent the most comprehensive treatment of the 
Myxomycetes in German since the work of Schinz in 1920. According to the authors, their 
work will help fill the gap since that time. In a special section dichotomous keys lead the 

a, with line drawings of habits on side by side pages next to the key. This 
idly key that should appeal to beginning students of the Myxomycetes . The 
es described as new: Cnbranafiltformh, C. macrospora, and C. stellifera. 
; prepared this work for professional botanists who may wish to identi 

)llecting and herbaria 

defined terms and 
e peoples names listed 

The author's classification system considers these organisms aligned with the fungi 
retaining the proper designated endings prescribed by the International Code of Botanical 
Nomenclature. The subordinal ranks follow closely the classification promoted by Nannenga- 
Bremekamp since many of her taxa, especially genera and families, are recognized. 

in the order Liceales. The order Liceales was a heterogenous unnatural a 



V convinci: 

ng t 

evidence to recogn 

ize twc 

) new families, t 



e, in the Liceales. Ir 

1 the ca 

se of Mmakatella, 


petty of bi 


ngence of spores an 

iitium, supposedly 

ilts in the 


laracter; a characte 


cies but IS I 


to move taxa from d 


orders and establi 


uencing tec 
er taxa sucl 


that has gone on in t 
ques to analyze gen, 
the protostelids and 

.t,c ref. 

omycetes should a' withm , 

The literature cited section is especially valuable since it contains much of th 
terature on the Myxomycetes since G.W. Martin and C.J. Alexopoulos pub 
?orld monograph, "The Myxomycetes," in 1969- This section is comprehensive 
itations from throughout the world. 

This book should be on the bookshelf of every student of the Myxomycetes. ' 
:iould consider an English translation to increase interest in the Myxomycetes o 

Sporobolus pojpsiensis (Poaceae: Eragrosteae): A new rhizomatous species from S 
Potosi, Mexico^and a new combination in S. airoides 
Joseph K. Wipffand Stanley D. Jones 

Notes on Carex, Cyperus, and Kyllinga (Cyperaceae) : 
species previously unreported to the state 
Charles T. Bryson and Richard Carter 



Validation of Kariuia (Cyperaceae) 207 

Penstemon //j//rte7(Scrophulariaceae) new to Texas 207 

Euphorbia i>raminea {¥.\\\)\\ovh'nce2it) new to Florida 208 

Melica subulata (l\)aceae: Meliceae): The first report for C 


Miconia skeaniana (Melastomataceae: I 
from eastern Cuba 
Walters Judd 

Cytological studies on North American species of 

(Poaceae: Andropogoneae) 

D. M. Burner and Roberto. Webster 

Systematic stud^ of Texas populations ot Phacelia p 


Jennifer Ann Moye) and BiUie L Turner 

William D Tidwell 


us Cybianthus subgenus Cybianthus ( 

A new species ot Carex (Cyperaceae Phaestoglochin ) from Oklahoma 
and Texas tvpihc ition ot section Phaestoglochin ind notes on sections 
Bracteosae and Phaestoglochin 



Lloyd H. Shinners 


Wm. F. Mahler 
Publisher 1971-1992 
Director Emeritus 

Botanical Research Institute of Texas, Ii 

509 Pecan Street 

Fort Worth, Texas 76102-4060 

H]7 332-4441/ 817 332-4112 FAX 

Dpto. de Botanica, Facultad de Biologia 
Universidad de Leon 
E-24071 Leon, Spain 




Department of Botany, 220 Bartram Hall 

University of Florida, Gainesville, PL 32611-8526; and 

Herbarium, 209 Rolfs Hall, Florida Museum of Natural History 

Gainesville, FL32611-0530, U.S.A. 

Mkonia skeaniana, which occurs in moist montane forests, cloud forests, and thickets from ca 
750-1974 m in the Sierra Maestra, is described and illustrated. It is compared with the closely 
related M. alternifolia, of the mountains of northern "Oriente," a species with which it has been 

In the course of taxonomic study of herbarium material in connection with a 
monograph of the Antillean members oiMiconia section Chaenopleura Benth. & 
Hook.f, the following undescribed species ofMiconia was recognized. Material 
representing this new species previously has been identified as M. alternifolia 
(Griseb.) Alain {=Miconia grisebachiiTn2SiQ). The new species clearly is referable 
to Miconia sect. Chaenopleura (see Cogniaux 1891), a distinctive, diverse, and 
presumably monophyletic group (at least within the Antilles) possessing an 
actinomorphic androecium (i.e., stamens forming a radially symmetrical pat- 
tern around style) of glabrous, white stamens of which the obovate anthers open 
by longitudinal slit-like pores (Judd & Beaman 1988; Judd & Skean 1991). The 
group is also characterized by an indumentum of usually ferrugineous, ± ir- 
regularly stellate- branched hairs, globose fruits that turn from red to pale blue 
at maturity, and angular-obovoid seeds with a + smooth testa (see Judd & Skean 
1 99 1 , fig. 1 OB). The section is especially well developed in the Greater Antilles, 
where many narrow endemics occur. The new species is described and compared 
with M. alternifolia, a species with which it has been consistently confused. 
Miconia alternifolia occurs in the mountains of northern "Oriente"^ from the 
Sierra de Cristal eastward, while M. skeaniana is restricted to the Sierra Maestra. 

rovinces of Cuba, i.e., Granma, Santiago de Cuba, 
which were created in the post- 1 959 pohtical reorganization of the 

"Oriente," the pre- 195 9 name of the political unit 

1 Force 


nience the four ea 



nd Guantanamo, % 






Miconia skeaniana Judd, sp. nov. (Fig. 1) 

Miconiae alternifoliae (Griseb.) Alain affinis, sed foliorum nervationibus magis valde abaxiale 

stellatis ca 0.09-0.2 mm latis in foliorum pagina abaxiali, interdum cum pilis minutis et 
mm longa, i.e., 66-77% antherae, (vs. 0.9-1.4 mm longa, i.e., 39-58% antherae). 

Shrub to 4 m tall. Indumentum of multicellular, fermgineous, minute-globu- 
lar, globular-stellate, or irregularly stellate-branched, to elongate-branched hairs. 
Young twigs not ridged, ± rectangular in cross-section, 1.7-4.5 mm wide, be- 
coming + terete with age, the indumentum of moderate to dense, irregularly 
stellate-branched to elongate-branched hairs, these ± persistent; internodes 1- 
4.5 cm long. Leaves opposite, with petiole 0.7-2.9(-3.2) cm long, the indumen- 
tum similar to that of the twigs; blade ovate to elliptic, 5-1 1 .2(-l6) X 1 .7- 
4. li-4A) cm, flat, coriaceous, the apex acuminate, the base acute to rounded or 
very slightly cordate, the margin plane to revolute, obscurely serrulate to undu- 
late-serrate, especially distally, proximal 0^0% of margin entire, but frequently 
appearing entire throughout (when revolute), the largest teeth to 0.2-0.4 mm 
(occasionally to 0.8 mm) long; venation acrodromous, slightly suprabasal to ± 
basal, with prominent midvein (primary vein) and 4 secondary veins, with 2 
conspicuous secondary veins placed 2-6.5 mm in from margin, 2 inconspicuous 
secondary veins placed closer to margin, and numerous percurrent tertiary veins 
oriented subperpendicular to midvein, the tertiary veins sometimes partially 
separated by composite inter- tertiary veins but usually connected by 1 to nu- 
merous quaternary veins, higher order veins ± orthogonal-reticulate; adaxial 
surface very soon glabrescent, the midvein and major secondary veins impressed, 
minor secondary veins, tertiary veins, and some quaternary veins slightly im- 
pressed, and higher order veins flat, the surface appearing minutely wrinkled- 
papillose after drying, with sparsely to densely scattered druse crystals; abaxial 
surface with moderate to dense, persistent, irregularly stellate-branched or globu- 
lar-stellate hairs, 0.09-0.2 mm across, on lamina and smaller veins, sometimes 
also with minute globular hairs, intergrading with slightly larger stellate- 
branched to elongate-branched hairs on midvein, the midvein and major sec- 
ondary veins prominently raised, tertiary veins prominently to slightly raised, 
minor secondary veins, some to all quaternary veins, and sometimes even a few 
higher order veins slightly raised. Inflorescences terminal, many-flowered, broadly- 
rounded cymes of 3 or 4 branch-pairs, 3-8 cm long, 3-8 mm across; proximal 
segment of lowermost inflorescence branches 1-2.9 cm long, distal internodes 
shorter, ultimate branches 1-6.5 mm long, with + moderate, irregularly stel- 
late-branched to globular-stellate or elongate-branched hairs; peduncle 1.5-5 
cm long, with similar indumentum; each branch associated with early caducous, 

skeaniana: A. flowering branch {Ekman 8855); B. leaf, abaxial surface (Ektnan 6928); C. detail of 
abaxial leaf surface (Ekman 6928); D. stellate hair from abaxial leaf surface {Seifriz 1072); E. 
flower (Ekman 8855); E stamen (Ekman 8855); G. young berry (Seifriz 1072). Mkonia altemifo- 
iia: H. leaf, abaxial surface (Ekman 6855); I. leaf, abaxial surface (Wright 1 79); J. detail of 
abaxial leaf surface (Shafer 8198); K. stellate hair from abaxial leaf surface (Ekman 6833); L. 
stamen, with elongate basal and sterile porrion of the anther (Ekman 6833). 

slightly ovate to obovate bract, ca 1 .7^ x 1-2 mm, the apex acute to rounded, 
the lowermost pair sometimes leaf-like; flowers in dichasia, usually distinctly 
separated from each other, each subtended by 2 caducous, ± ovate bracteoles, 
1.5-2.2 X 0.8-1 mm, the apex acute to obtuse; pedicel 0-0.5 mm long. Hj- 
panthium cylindrical, free portion 1 .6-1 .9 mm long, the outer surface with sparse 
to moderate, minute, stellate-branched hairs, the inner surface glabrous and 
strongly ridged, i.e., with 10 prominent ridges alternating with 10 weaker 
ridges, the apices of the stronger ridges projecting, 0.05-0. 1 mm. External calyx 
lobes (=teeth) 5, 0.3-0.8 x 1 .4-2.5 mm, broadly triangular, with acute to acumi- 
nate apex, indumentum as on hypanthium;i«/er;z^/a2/)/x/o^ej 5, 0.5-1.5 X 1.4- 
2.5 mm, ± triangular to ovate-triangular, green, glabrous or with sparse stellate 
hairs, apex rounded, margin entire, sometimes minutely fringed; calyx tube 
0.2-0.5 mm long. Petals 5, broadly ovate to obovate, 2.8-4.2 x 2.6-2.9 mm, 
glabrous, white to pink-tinged, imbricate and apically interlocking in bud, 
with apex emarginate, with an asymmetrically located notch; margin entire. 
Stamens 10, geniculate, glabrous; proximal segment (filament) 1 .9-2.9 mm long; 
distal segment (anther and connective) 2.6-3.2 mm long, with minute dorsal 
projection (pointing ± toward anther apex), the anther 1.8-2.5 mm long, with 
fertile portion (anther sacs) 1.2-1 .9 mm long, opening by 2 longitudinal slits, 
the connective extending 0.6-0.9 mm beyond the base of the anther. Ovary (2- 
or) 3-loculate(N=l, 10), ca 1/2-inferior (immature) to ca2/3-mferior (mature), 
short-ovoid to subglobose, 2.1-3 x 2.4-3.5 mm, glabrous and strongly ridged, 
with fluted apical projection to ca 0.5 mm encircling the base of style; style 3.8- 
5 mm long, terete, glabrous; stigma truncate. Bernes globose to subglobose, ca 
4.5-7 X 5.5-7 mm, pale blue. Seeds angular-obovoid, ca 0.5-0.9 mm long; 
testa smooth. 

Type: CUBA. Prov. Santiago de Cuba [=Oriente, p.p.}: Sierra Maestra, Cordillera de la 
Gran Piedra, La Gran Piedra, cloud forest, ca 1200 m ale, 10 Nov 1917, E.L. Ekman 8833 

Etymology. It is a pleasure to name this distinctive species after Dr. James Dan 
Skean, Jr. (b. 1958), plant systematist at the Department of Biology, Albion 
College, Michigan, who has assisted the author during fieldwork conducted in 
connection with a taxonomic revision of the Antillean species o( Mkonia sect. 

Additional Specimens Examined: CUBA. Prov. Granma [ = Oriente, p.p.}: Sierra Maestra, 
La Bayamesa, on the ridge between Rio Oro and Rio Yao, 1 100-1400 m, Ekman 7213 (F); peak 
of Punta de Palma Mocha, south of Yara, 1400 m, Ekman 14317 (NY). Prov. Santiago de 
Cuba [=Oriente, p.p.}; Sierra Maestra, summit of Pico Turquino, I960 m, Aama 6760 (NY); 
Pico Turquino, northern slopes, ca 1750 m, Ekman 3273 (S); ibid., Ekman 3389 (S); between 
Finca Reunion and peak of Loma del Gato, 750 m, Ekman 6928 (NY); near summit of Pico 
Turquino, Leon 10744 (GH, NY); between the arroyos Peladero and Indio, 3000-4500 ft, 
Lopez-Figmtras 406 (US); Gran Piedra, 1250 m, L6pez-F tgueiras 2661 (IJ, US); Pico Turquino, 

, Miconiaskeaniana 





20 y 

f£ ^ • 







J i072 (US); Gran Piedra,( 

Mkonia skeaniana is endemic to Cuba and known from several localities in 
the Sierra Maestra, both the Cordillera de Turquino and the Cordillera de la 
Gran Piedra, of southern "Oriente" {prov. Granma and Santiago de Cuba] (Fig. 
2), where it occurs in moist montane forests, cloud forests and thickets from 
750-1974 m elev. The vegetation of the higher elevations of the Sierra Maestra 
is summarized in Leon (1924, 1946), Seifriz (1943), and Borhidi (1991). 

Mkonia skeaniana is probably most closely related to M. alternifolia, with 
which it consistently has been confused. It can be readily distinguished from 
this species by its more strongly abaxially raised-reticulate leaf venation, i.e., 
midvein and major secondary veins prominently raised, tertiary veins slightly to 
prominently raised, minor secondary veins, some to all quaternary veins, and sometimes 
even a few higher order veins slightly raised {ws. midvein and major secondary veins 
prominently raised, minor secondary veins slightly raised, tertiary veins slightly 
raised to ±flat, and higher order veins flat). The leaves of M. skeaniana have consis- 
tently acuminate apices while those of M. alternifolia vary from broadly obtuse 
to acute or acuminate. Miconia skeaniana possesses a ferrugineous indumentum 
of irregularly stellate-branched to occasionally globular-stellate hairs, ca 0.09- 
0.2 mm across, on minor veins and lamina of the abaxial leaf surface. These hairs 
are sometimes intermixed with minute-globular hairs. In contrast, the hairs of 
M. alternifolia are more darkly ferrugineous, consistently globular-stellate, and 
smaller (i.e., 0.04-0.09 mm across). The basal and sterile portion of the anthers 

230 SiDA 16(2) 1994 

of M. skeamana is shorter than that in M. alternifolia, i.e., anther with fertile 
portion 1.2-1.9 mm long and occupying GG-11% of anther length in M. 
skeamana (vs. 0.9-1 .4 mm long and occupying 39-58% of anther length in M. 
alternifolia). Additionally, the tertiary veins of M. skeaniana are frequently more 
closely spaced than those of M. alternifolia, i.e., 5-12 (avg. 9) veins vs. 4-1 1 
(avg. 7) per 2 cm in middle portion of leaf. The two species are completely 
allopatric and are, thus, geographically isolated. 

The type oiMiconia alternifolia, collected by Charles Wright (no. 1 79), is 
actually a mixed collection composed of material collected at Monte Verde (Prov. 
Guantanamo) on March 2nd, 1859 (see original label on isotype at GH and 
label on holotype at GOET) and presumably at Loma del Gato in the Sierra 
Maestra (Prov. Santiago de Cuba) in 185(^1857. The exact locality of the 185<^ 
1857 collections is not known because no original label is present; however, 
plants matching these specimens occur only in the Sierra Maestra and it is known 
that Wright did collect in the Loma del Gato area of the Sierra Maestra in 
1856-1857 (Underwood 1905; Howard 1988). These collections of Charles 
Wright were all assigned the same number by Asa Gray (Howard, 1988). The 
holotype oi Mkonia alternifolia (at GOET) represents the "Monte Verde ele- 
ment" (collected on March 2,1859), which is considered to be conspecific with 
phenetically similar collections from other localities in northern "Oriente," i.e., 
prov. Holguin and Guantanamo; isotypes representing this collection are found 
at BM, GH, NY, and S (all with printed labels giving the incorrect date of 
1860-1864). Collections oi Wright 1 79 at GH, GOET, and M (collected in 
1856-1857, probably at Loma del Gato) actually represent the taxon here con- 
sidered to be M. skeaniana. As discussed above, this species is restricted to the 
Sierra Maestra. The name M. alternifolia is, of course, linked to the holotype 
specimen oiWright 1 79 (which represents the northern "Oriente" taxon). Thus, 
the plants of the Sierra Maestra, if considered specifically distinct from M. al- 
ternifolia, are left without a name, and herein are described as M. skeaniana. 

The specimens oiWright 1 79 collected in 1856-1857 have sometimes been 
considered as a distinct variety, i.e., M. grisebachii var. reticulata Cogn., but the 
type of this name actually is a Puerto Rican plant of the Sierra de Luquillo (and 
this name is a synonym of M. pychoneura Urb.). 

The recognition oiMiconia skeaniana brings the number of Antillean species 
ofMiconia sect. Chaenopleura to 43 (Judd, unpublished data). Other members of 
this section occurring in Cuba include: M. alternifolia, M. cubensis (Griseb.) C. 
Wright, M. rufa (Griseb.) Triana (probably conspecific with M . plumeriaefolia 
Britton & P. Wilson), and M.. subcorymbosa Britton (probably conspecific with M. 
calycina Cogn.). 

lank Wendy B. Zomlefer for : 

BORHIDI, A. 1991 . Phytogeography and vegetation ecology of Cuba. Akademiai Kiado, Budapest. 
CcxiNiAUX, A. 1981. Melastomaceae. In: A. C. de CandoUe, editors. Monographiae 

Howard, R.A. 1988. Charles Wright in Cuba, 1856-1867. Chadwick-Healey, Alexandria, 

JuDD, W.S. and R.S. Beaman. 1988. Taxonomic studies in the Miconieae (Melastomataceae). II. 

Systematics of the Miconia sukompressa complex of Hispaniola, including the description of 

two new species. Brittonia 40:368-391. 

IV. Generic realignments among terminal-flowered taxa. Bull. Florida Mus. Nat. Hist., Biol. 
Le6n, Hno. 1924. Una excursion al Pico Turquino. Mem. Soc. Cub. Hisr. Nat. "Felipe Poey" 
Le6n, Hno. 1946. Flora de Cuba. Vol. 1 . Contr. Ocas. Mus. Hist. Nat. Colegio "De La Salle" 

c. Club 32:291-31 


Kartesz, John. T. 1994. A Synonymized Checklist of the Vascular Flora 
of the United States,Canada and Greenland. Second Edition. (ISBN 
0-88192-204-8, hbk, 2 volumes); Timber Press Inc., 9999 S. W. 
Wilshire, Suite 124, Portland, OR 97225. (800) 327-5680. $159.00. 
Vol. 1,622, vol 2, 816 pp. 

This is officially a second edition, but it is more than just a new edition, it is father a much 
expanded major rework which includes a new format and a second volume. This is a significant 
improvement over his 1980 work, which has been the standard for fourteen years. 

As monographic research continues the taxonomic and nomenclatural process is in constant 
flux, and will be for many years to come. Also, much of the classification of our flora is a tangle 
of individual taxonomic opinion. This work does not attempt to solve all these problems, but it 
is a quantum leap forward, and is a solid new foundation on which to build the next cycle. With 
Kartesz' new Checklist we now have an incredibly valuable and comprehensive effort that sets a 
new standard. 

The new edition has two volumes. The first is the familiar checklist that is alphabetical by 

actiy those in the body of the checklist with each name listed alphabetically by genus. Accepted 
names appear in Roman print and stand alone, whereas synonyms appear in italics and are 
followed by their accepted names. The thesaurus differs from the checklist by offering a rapid 
location system for all names, save those names above the genus level. The thesaurus and the 
checklist are together intended to provide a comprehensive summary of accepted names and 
their synonyms of the North American vascular flora north of Mexico: the continental United 
States, Hawaii, Puerto Rico, and the U. S. Virgin Islands; Canada; the French islands of St. 
Pierre and Miquelon; and Greenland. 

This is a major work which is used many times a day in my work of compiling floras, 
manuals, articles, books, and plant checklists of Texas and i 
Checklist on plant nomenclature and its synonymy is 





United States Department of Agriculture 
Agricultural Research Service 

Sugarcane Research Unit 

Houma, LA 70361, U.S.A. 


United States Department of Agriculture 

Agricultural Research Service 

Systematic Botany & Mycology Laboratory 

2n = 60), S. coarctatum (2n = 60), and S. giganteum (2n = 30 
the first report of a polyploid series within S. giganteum and t 
tnthus group. Counts also were made for S. alopecuroideum (2: 

tccharum L. (incluyendo Erianthus Michx.) na 
rizada. Nuestros objetivos fueron determii 

5 de Saccharum spp.). Se ofrecen por primera vi 
I, y 5. coarctatum (2n = 60), y .S. giganteum (2n = 
ta de una serie poliploide en S. giganteum y el p 


There has been disagreement among taxonomist 
and placement oi Saccharum L. and Erianthus Michx 

OLis concepts of these genera is presented in Webster & Shaw (1995). North 
American taxonomists have followed the concepts of Hitchcock (195 1) and 
have recognized both genera. European agrostologists have placed Erianthus 
within Saccharum. Characteristics used to distinguish the genera are difficult to 
apply when the full range of variation is considered. Therefore, we are following 
the concept presented in Clayton & Renvoize (1986) that recognizes Erianthus 
as a synonym o'i Saccharum. 

Sugarcane breeders recognize separate genera and generally include five to six 
species within Saccharum: S. edule Hassk. (2n ^ 60 to 80), S. offictnarum L. (2n = 
80), S. rohustum Brandes & Jesw. ex Grassl (2n = 60, 80, to about 200), S. sinense 
Roxb. (2n = 1 1 1 to 1 20), and S. spontaneum L. (2n = 40 to 1 28). A sixth species, 
S. barberijesw. (2n = 81 to 124), is sometimes included in^. smense. Chromo- 
some counts (Daniels et al. 1975) for closely related taxa called the "Saccharum 
complex" by Mukerjee (1957) include Old World Enanthm Michx. sect. Rip- 
idium Henrard (2n = 20, 30, 40, 60); Miscanthus Anderss. sect. Dtandra Keng 
(2n = 40); Narmga Bor (2n = 30); and Sclerostachya (Hack.) A. Camus (2n = 30). 
These and a few other genera have at times been placed in Saccharum (Daniels 
and Roach 1987; Whalen 1991). The basic genomes within Saccharum (s. str.) 
appear to be x = 8, 10, and 12 (Sreenivasan et al. 1987), and that olEnanthus 
may be x = 5, typical of the Andropogoneae (Celarier 1956a). 

Harlan and de Wet (1975) noted that Saccharum had an "oversplit taxonomy," 
implying that many taxonomic divisions may be artificial. As evidence of this, 
interspecific and intergeneric crosses within xhtSaccharum complex are usually 
successful (Gill and Grassl 1986; Grassl 1980; Daniels and Roach 1987). The 
taxonomic relationships among the taxa of the Saccharum complex have been 
neither carefully studied nor well-defined (Webster and Shaw 1995) and they 
conclude that Erianthus is best treated as a synonym oiSaccharum. Five species 
and one variety oi Saccharum L. were recognized by Webster and Shaw (1995) as 
being native to North America. They are 5". alopecuroideum (L.) Nutt. {= Erian- 
thus alopecuroides (L.)^\\.'\,S. haldwinii Spreng. (= E. strictusBa.ld'w.),S. brevibarbe 
(Michx.) Pets. var. brevibarbe, S. brevibarbe (Michx.) Pers. var. contortum (Nutt.) R. 
Webster (= E. contortus Ell.), S. coarctatum Fern. (= E. coarctatus Fern.), and S. 
giganteum (Wait.) Pers. {= E. giganteus (Walt.) C.E. Hubb.}. 

Old World species previously treated in Erianthus (sect. Ripidium) are used 
in sugarcane breeding (Heinz 1991), particularly for its disease resistance (Burner 
et al. 1 993 ; Grisham et al. 1 992) and freeze tolerance (Moore 1 987). Chromo- 
some numbers of many clones have been reported (Babu and Srinivasan 1960; 
Mohan and Sreenivasan 1983). Hybrids between Saccharum spp. and North 
American species placed in Erianthus have not been reported, although Gill and 
Grassl (1986) reported hybrids between Sclerostachya fusca (Roxb.) A. Camus 
and E. brevibarbis Michx. (= S. brevibarbe), E. tracyi Nash. (= S. alopecuroideum), E. 

Webster, Cytological studies on Saccharum 235 

contortus Baldw., and between Mtscanthus sp. and E. brevibarbis Michx. (= S. 

Except for the few clones (five clones in four species) reported by Celarier 
(1956b) and Gill and Grassl (1986), there have been no cytological studies of 
the North American species traditionally placed in Erianthus. The objectives of 
this research were to determine chromosome numbers and characteristics in a 
complete collection of North American species of these taxa and attempt hy- 
bridization with elite sugarcane hybrids (interspecific and mtetg^enenc Saccharum 
spp. hybrids). 


Clones were collected as rhizomes from natural populations in the mid-At- 
lantic and southeastern U.S.A. (Fall 1992) and southcentral U.S.A. (Fall 1993) 
and were grown at Houma, Louisiana (29° 35'N 90° 44' W) in 18.9 L cans filled 
with a soil mix of 3:2:2 (soil:sand:peat moss). Taxonomy was verified according 
to the concepts presented in Webster and Shaw (1995). Voucher specimens are 
deposited at TAES. 

Inflorescences were collected at early boot stage and fixed in Carnoy's B: etha- 
nol, acetic acid, and chloroform (6:3:1 mixture by volume) (Smith 1947) and a 
few drops of saturated ferric chloride. Anthers were squashed in 5 g L"^ propi- 
onocarmine. Chromosome number was determined from pairing configurations 
at diakinesis or metaphase I (MI) in 5 to 25 microsporocytes per plant. Chromo 
some number for some plants was determined or verified in squashes of root tij 
cells using standard procedures. Pollen stainability, an estimate of pollen viabil 
ity, was measured by staining mature anthers in 10 g L"^ I2-KI. 

The cross-sectional area of individual bivalent chromosomes was measured a 
MI for each of nine clones (range 1 to 1 5 cells clone"^ mean 6 cells clone"^) using 
a Cue-2' image analyzer (Galai 1990). Area (pm^) was determined from the 
number of pixels in the bivalent chromosome. 

About 100 completely filled pollen grains from mature anthers of 25 plants 
were collected in Fall 1993, stained in 10 g U^ I2-KI, and imaged at 400x. 
Cross-sectional area was calculated as described above. Average cross-sectional 
radius (pm) of each pollen grain was the mean of eight Martin's radii measured 
at 0,45, 90, 135, 180, 225, 270, and 315° (Galai 1990). Volume (pm^) was 
calcu-lated from average radius assuming that pollen grains were perfect spheres. 

Analysis of variance of pollen area and volume was done by the general linear 
models procedure (SAS Institute 1990). Sources of variation were species (7 df), 

clone within species (16 dO, and residual (2,773 dO- The effect of species was 
tested using clone within species as error; clone within species was tested using 
residual as error. Means were compared using least significant difference at the 
5% level of significance (Steel and Torrie 1980). The cluster procedure (SAS 
Institute 1990) was used to assign clones to non-overlapping hierarchical groups 
based on multivariate analysis of chromosome number, pollen area, and pollen 
volume. Prior to conducting the cluster analysis, data were standardized to a 
mean of and standard deviation of 1 using procedure "standard" (SAS Insti- 
tute 1990). 

Crosses between the native North American taxa oiSaccharum and elite sug- 
arcane clones, cytoplasmically male-sterile (cms) Sorghum bkolor(L.) Moench, 
and Vetiveria zizanioides (L.) Nash (used as females) were attempted in Fall 1 993 . 
Flowering of sugarcane clones was induced using standard photoperiod treat- 
ment (Dunckelman and Legendre 1982). Flowering o^ Sorghum, North Ameri- 
can 5'<^fr^(:?r?i(w, and V. zizanioides occurred under natural photoperiod. Conven- 
tional methods of crossing, seed maturation, seed germination, and seedling 
establishment were used (Dunckelman and Legendre 1982). 


Chromosome number varied among and within the North America Saccharum 
species (Table 1). Saccharum alopecurotckum and S. haldwinii were 2n = 30. Counts 
of 2n = 30 in E. stnctus from Texas (Celarier 1956b) and E. tracyi (Gill and Grassl 
1986) are consistent with our data. A count of 2n = 60 in S. alopecuroideum 
(Celarier 1956b) is inconsistent with our data. However, two clones of 5'. gigan- 
teum (2602 and 2603) v^ith 2n = 60 and characteristics similar to S. alopecuroi- 
deum were collected in Tennessee and Arkansas. It seems probable that Celarier's 
count of 2n = 60 may be based on S. giganteum according to present taxonomic 
concepts. Clones with 2n = 30 and 2n = 60 are probably 6x and 12x, respectively. 

Saccharum giganteum consisted of clones with 2n = 30, 60, and 90 chromo- 
somes. These are the first counts for the species, the first indication that the 
species is a polyploid series, and the first report of 2n = 90 (18x) in the taxa 
traditionally placed in Erianthus. The 2n = 30 types were collected from Mary- 
land south to Alabama and in southern Louisiana; the 2n = 60 types were col- 
lected in Alabama, Arkansas, Georgia, Louisiana, and Tennessee; and the 2n - 
90 types were collected in southern Georgia and Florida. This indicates a geo- 
graphic effect on distribution of cytotypes. Future studies are planned to define 
the relationship between chromosome number and morphology within this 
species. Saccharum bengaleme Retz. 1= E. bengalense (Retz.) Bharadw.} has also 
been shown to be a polyploid series with 2n = 20, 40, and 60 c 
(Mehraetal. 1968). 

\BAMA. Blount Co.: Hwy 278, 32 km W of Gadsden, 6 Nov 1992, Webster 2551 (TABS). 
mar Co.: Hwy. 18, 24 km E of Fayettsville, 6 Nov 1992, Webster 2548 (TAES). GEORGIA, 
nks Co.: Interstate 85, 3 km E of Commerce, 7 Nov 1992, Webster 2553 (TAES). Brooks 
.: Hwy 84, 8 km E of county road 219, 27 Oct 1992, Webster 2533 (TAES). Forsyth Co.: 

. Perry Co.: Hwy 10, 2 km E of Hwy 

I, 27 Oct 1992, Webster 2532 (TAES). 

y 82, 2 km E of county road 29, 28 Oct 1992, Webster 2538 
1 km S of county road 65, 28 Oct 1992, Webster 2539 (TAES). 
ite line, 27 Oct 1992, Webster 2536 (TAES). Lamar Co.: Hwy 
fov 1992, Webster 2547 (TAES). Pickens Co.: Hwy 82, 13 km 
:, Webster 2542 (TAES). ARKANSAS. White Co.: Hwy 6A, 6 
Webster 2606 (TAES). Yell Co.: Hwy 10, 2 km W of Birta, 10 
). GEORGIA. Decatur Co.: Hwy 84, 1 km from jet 285, 14 

Hwy 13, S of Salisbury, 23 Oct 1992, Webster2502 (TAES); Hwy 13, 1 1 km N ( 
Oct 1992, Webster 2503 (TAES). MISSISSIPPI. Choctaw Co.: Natchez Trace I 
Hwy 9, 28 Oct 1992, Webster 2543 (TAES). NORTH CAROLINA. Greene C 
km S of Greenville, 24 Oct 1992, Webster 2509 (TAES). Halifax Co.: Hwy 
Hwy 97, 24 Oct 1992, Webster 2508 (TAES). SOUTH CAROLINA. Florence 
3 km N of Langston Rd, 25 Oct 1992, Webster 2514 (TAES). TEXAS. Angelir 
6 km N of 7 jet., 13 Nov 93, Webster 2611 (TAES). Cherokee Co.: Hwy 1 
Jacksonville, 13 Nov 93, Webster 2609 (TAES). VIRGINIA. Accomack Co.: H 
of Parkway, 2 km N of Keller, 23 Oct 1992, Webster 2504 (TAES). 

km N of state line, 27 Oct 1992, Webster 2531 {TKE^). 
km N of Starke, 26 Oct 1992, Webster 2521 (TAES). 
.4 km N of Nahunta, 26 Oct 1992, Webster 2523 (TAES). 
ochee River, 25 Oct 1992, Webster 2520 (TAES). Grady 
1992, Webster2534 (TAES). Screven Co.: Hwy 301, 8 
r 2579 (TAES). Wayne Co.: Hwy 301 , 3 km N of Jones 

4 Oct 1992, Webster 25 13 (TAES). 

5 Oct 1992, Webster 2516 (TAES). 

NORTH CAROLINA. Robeson Co.: Hwy 301, 5 km N of Lit 
Webster 2512 (TABS). Wayne Co.: Hwy 13, 2 km N of Racl 
Webster 2510 (TABS). MARYLAND. Wicomico Co.: Hwy 
1992, Webster 2501 (TABS). SOUTH CAROLINA. Bamb( 


Oct 1% 




ccomack Co.: 



)2, Weki 


pton Co.: Hwy 258, 3 km N of NC state 


e',24 0c 


2, Wehst 

er 2506 (TABS). 

2n = 60- AL 


\ Cu 

llman < 

:o.: Hwy 91, at 4 I 

Tiile mai 

rker, 6 Nov 1' 

)92, Webster 2550 (TAES). 





Webster 2546 { 


HS). ARKANSAS. White Co.: Hwy 367, 




Umks, 9 nIv 199' 


). GBORGIA. Evans Co.: 

)ct 1992 




9, 53 km N of Opel 



[wy70A bypass, Nc 

ngton, 8 Nov 1 

1993, Webster 2602 (TAES). 

?n = 90FK 



L- Co.: I ; 

Iwy 19&98 2 km; 

5 of coun 




\ES) Ty 


,vy51 10 km from 

the Gul 

f of Mexico, 2 


NofPerry,27 0ct] 


ES). GEORGIA. Charlton 




2526 (TABS). 

Saccharum brevibarbe-v2i.K. brevibarbe and contortum a 
60. Gill and Grassl (1986) reported that a clone of £, brevtbarbh was 2n = 60, 
which confirmed our finding for the species. However, they reported that a 
clone of £. contortus was 2n = 30, which may be the result of misidentification. 
Saccharum brevibarbe var. contortum is characterized as having callus hairs equal to 
or shorter than the spikelet, while callus hairs are either absent {S. baldwinii) or 
longer {S. giganteum and S. alopecuroideum) than the spikelet in 2n = 30 types 
(Webster and Shaw 1995). 

Meiosis was normal in all clones and univalents and quadrivalents were ex- 
tremely rare (Fig. 1). Celarier (1956b) noted bivalent size polymorphism m 
North American Saccharum. Mean bivalent area of 4.43 pm- (range 1 .58 to 8.94 
pm^), 3.25 pm^ (range 1.10 to 8.77 pm^), and 4.43 pm^ (range 1.86 to 7.6lpm^) 
in 2n = 30, 60, and 90 biotypes, respectively, differed little among cytotypes. 
Bivalent area was not normally distributed; the distribution was generally shifted 
toward the smaller size classes. Bivalents of an Old World clone (£. rufipilus, 2n 
= 20) averaged 3.89 pm- (range 3-25 to 5.28 pm-), and those of an elite sugar- 
cane clone LCP 81-30 (2n = 105, 108) averaged 3.42 pm^ (range 1.55 to 
6.99 pm^) (Burner and Legendre 1994; Burner unpublished data). Thus, 
North American Saccharum tends to have wider ranges of bivalent area than 
Old World Saccharum, but there is little difference in area or range between 
New World Saccharum and sugarcane. The data support the observation by 
Gould (1956) that there is no obvious correlation between chromosome 
number and chromosome size in the Andropogoneae. 

Webster, Cytological studies on Saccl 



• * 


•** . • 



B / • 



^ ♦ 

<* 1- 

« • 


-w- ... 


# • 












^ ♦ 

» — 

MG. 1 . Meiotic metaphase 1 chromosomes of North American Sacchamm. Bar i 
epresencs 10 \im. {A)S. gtgantmm (n = 15 bivalents) [x 1600]. (B)5. brevibarbev^ 
= 30 bivalents) [x 1600]. (C) 5. gigantmm (n = 45 bivalents) [x 1000}. 

Pollen area and volume differed significantly among species and clones 
within species (Table 2). As expected, species with 2n = 30 had smaller 
pollen than those with 2n = 60 or 2n = 90 (Gould 1957). Pollen size has 
been used as a taxo-nomic character to separate closely related taxa differing 
in chromosome number (Gould 1953 & 1957). However, variation in pol- 
len size among clones within species indicates that pollen size would be an 
unreliable predictor of species. 

Clones were assigned to clusters based on chromosome number, pollen area, 
and pollen volume. The dendrogram beginning with seven clusters (R^ = 0.97) 
and ending with one cluster (R^ = 0.00) is shown in Fig. 2. There was a 


S. giganteum 



S. giganteum 
S. giganteum 







s'. hrJiiteSe var. 



Fig, 2. Dendrograrr 





Volume (pm3) 





S. baldtvinii 





ir, brev 








LSD (0.05) -Clont 

10% loss in explained variance when seven clusters were combined to four 
clusters. Four-cluster analysis (R- = 0.87) seemed to be most informative. 
Clones ofS. giganteum (2n = 60) and Webster 2316 {S. coarctatum, 2n = 60) 
were assigned to cluster group 1 . Morphology of Webster 2516 is otherwise 
typical o'lS. coarctatum. Clones of 5, giganteum with 2n = 90 chromosomes 
were assigned to cluster group 2. Clones of 5", brevibarbe vars. brevibarbe and 
contortum and S. coarctatum, except for Webster 2316 were assigned to cluster 
group 3. Clones with 2n = 30 were assigned to cluster group 4. Thus, the 
four-cluster analysis was generally consistent with present taxonomic con- 
cepts (Webster and Shaw 1995) and provided evidence of diversity be- 
tween the cytotypes of S. giganteum. There was a further loss of 11% of 
explained variance when the 2n = 60 and 2n = 90 cytotypes oiS. giganteum 
were joined to form three clusters. The three-cluster analysis explained 
76% of variance. Only 50% of total variance was explamed by two clusters. 
Crosses were successfol between elite sugarcane and North American Saccharum 
(Table 3). A cross with Webster 235 1 (2n = 90) yielded 448 seeds, and other crosses 
yielded to 92 seeds. The potential agronomic value of these Fj hybrids will be 
evaluated in subsequent tests. Two crosses of cms Sorghum x North American 
Saccharum (five crosses attempted) yielded some seed, but grow-out evaluation 
showed that the Fi progeny were not hybrid. Crosses of cms Sorghum x elite 
sugarcane also failed to yield hybrid progeny (Burner unpublished data). Vetiveria 
ztzantotdes (2n = 20) X Webster 2352 (2n = 30) was also unsuccessful. Pistil- 
late sterility is frequently observed in V. zizanioides (Ramanujam and Kumar 

1963), and despite several attempts we have never obtained viable seeds from 

Love (195 1) noted that differing chromosome numbers within a species, 
as we found in S. giganteum, indicates the species may include more than 
one distinguishable taxon and needs closer taxonomic inspection. Love 
(1951) and Nannfeldt (1938) further argue that intraspecific difference in 
ploidy level has fundamental systematic value sufficient to justify the rec- 
ognition of their respec-tive members as species. Cytomorphological study 
of 5", gigantemn should be conducted and crosses between 6x and 18x forms, 
to attempt synthesis of the intermediate \2x form, would further our un- 
derstanding of the evolution of the genus. 

We thank B. Duet and S. Michalisko, USDA-ARS, Houma for t 
assistance, and Drs. P. Bramel-Cox (Manhattan, Kansas) and F. MiJ 
lege Station, Texas) for providing sorghum seed. 

Babu, C.N. and K. Srinivasan. I960. Chromosome numbers in the genus Erianthus. Sci. 

Cult. 26:230-231. 
Burner, D.M., M.P. Grisham, and B.L. Legendre. 1993. Resistance of sugarcane relatives 

injected with Usttlago Kitamtnae. Plant Dis. 77:1221-1223. 
Burner, D.M. and B.L. Legendre. 1994. Cytogenetic and fertility characteristics of eLte 

sugarcane clones. Sugar Cane 1994 (1):6-10. 
Celarier, R.P. 1956a. Addirional evidence for five as a basic number of the Andropogoneae. 

Rhodora 58:135-143. 

Saccharinae. Cytologia 21:272-291. 
Clayton, W.D. and S.A. Renvoize. 1986. Genera graminum: Grasses of the World. Kew, 

London: Royal Botanical Gardens. 
Daniel.s,J., p. Smith, N. Paton, and C.A. Williams. 1975. The origin of the genus ^^ir^^raM. 

Sugarcane Breed. News!. 36:24-39. 
Daniels, J. and B.T. Roach. 1987. Taxonomy and evolution, pp. 7-84. In: D.J. Heinz 

Dunckelman, P.H. and B.L. Legendre. 1982. Guide to sugarcane breeding in the temper- 
ate zone. USDA-ARS ARM-S-22. Agric. Res. Serv. Southern Region. USDA, New Or- 
leans, Louisiana. 

Olympus Corp., Lake Success, New York. 
Gill, B.S. and CO. Grassl. 1986. Pathways of genetic transfer in intergeneric hybrids of 
sugar cane. Sugar Cane 1986(2):2-7. 

244 SiDA 16(2) 1994 

1957. Pollen size as related to polyploidy and speciation in the Andropogori 

saccharoides-A. barbinodts complex. Brittonia 9:71-75. 
Grassl, CO. 1980. Breeding Andropogoneae at the generic level for biomass. Sugarcane Breed. 

Grisham, M.R, D.M. Burner, and B.L. Legendre. 1992. Resistance to the H strain of 

Harlan, J. R. and J.M.J, de Wet. 1975. On O Winge and a prayer: The origins of polyploidy. 

Bot. Rev. 41:361-390. 
Heinz, D.J. 1991. Sugarcane cytogenetics, pp. 279-293. In: T. Tsuchiyaand P.K. Gupta (eds.) 

Chromosome engineering in plants: Genetics, Breeding, Evolution, Part B. Elsevier, 

Hitchcock, A.S. 1951. Manual of the grasses of the United States, 2nd ed. (revised by A. 

Chase). U.S. Dept. Agric, Misc. Publ. No. 200, Washington, D.C. 
Love, A. 195 1. Taxonomical evaluation of polyploids. Caryologia 3:263-284. 
Mehra, RN., PK. Khosla, B.L. Kohli, and J.S. Koonar. 1968. Cytological studies in the 

North Indian grasses (part 1). Res. Bull. Punjab. Univ. 19:157-230. 
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Erianthus. Sugarcane Breed. Newsl. 45:50-63. 
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improvement through breeding. Elsevier, New York. 
MuKHERjEE, S.K. 1957. Origin and distribution oiSaccharum. Bot. Gaz. 1 19:55-61 . 
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tetraploids of sect. Ochlopoa A. & Gr., and some additional notes on Ochlopoa. Svensk. Bot. 

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Ramanujam, S. and S. Kumar. 1963. Irregular meiosis associated with pollen sterility in Vetweria 

Sas Institute. 1990. SAS/STAT users guide, version 6, fourth ed. SAS Institute, Gary, North 

Smith,L. 1947. The acetocarmine smear technic. Stain Technol. 22:17-31. 
Sreenivasan, TV., B.S. Ahloowalia, and D.J. Heinz. 1987. Cytogenetics, pp. 211-253. In 

D.J. Heinz (ed.) Sugarcane improvemenr through breeding. Elsevier, New York. 
Steel, R.G.D. and J.H. Torrie. 1980. Principles and procedures of statistics. McGraw-Hill, 

New York. 
Webster, R.D. and R.B. Shaw. 1995. Taxonomy of the native North American species of 

Saccharum (Poncea.e: Andropogoneae). Ann. Missouri. Bot. Gard. (submitted) 
Whalen, M.D. 1991. Taxonomy ofSaccharum (Poaceae). Baileya 23:109-125. 



Department of Botany 

The University of Texas 

Austin, TX 787 13, U.S.A. 

Populations of Phace/ia patuliflora 
irieties, P patulifora y^T. patuliflora a 

, a species of the southeastern United States and closely 
n Texas. The species was formerly treated as having two 
nd P. patuliflora var. teucriifolia. The present investigation 
ariety from south Texas be recognized for which the name 

patuliflora var. austrotexana]. A. 
)mplex along with a map showing t 

Moyer, var. nov. is proposed. A key to the varieties of this 

Seinvestigaron en Texas poblacion 
)s Unidos y partes adyacentes de Me 
fora var. patuliflora y P patuliflora vai 
r reconocida una vanedad regional a. 
patuliflora. ^r.austrotexana].h.m 

ies Phacelia patuliflora, una especie del sureste de los Esta- 
xico. Se reconocian anteriormente dos variedades, P. patu- 
: teucriifolia. La presente investigacion sugiere que puede 
dicional del sur de Texas, para la que se propone el nombre 
jyer, var. nov. Se incluye una clave para las variedades de 

Key Words: Hydrophyllaceae, Phacelia, Texas 

Phacelia patuliflora belongs to section Cosmanthus Brand of the Hydrophyl- 
laceae. This section is characterized by the presence of a gland or nectary on the 
principal vein of the corolla (Constance 1949). Cosmanthus consists of 14 species 
with a south to north distribution starting in the Guatemalan and the Mexican 
highlands and extending northwards into Texas and Oklahoma and eastward 
through the Ozarks to the Appalachians. Other species oi Phacelia do not over- 
lap the distribution of the subgenus Cosmanthus, with the exception of P. congesta 
Hook, and its few relatives, which are not morphologically or cytologically similar 
to Cosmanthus (Constance 1949). Gillett (1968), in his cytological work on Cos- 
manthus, also recognized 14 species in the section. Turner (1991) has recently 
added three additional new species, P. altotonga, P. carmenensis, and P. neffli, to the 
Cosmanthus complex. 

The Phacelia patuliflora complex is most commonly aligned with P. strictiflora 
and P. laxa (Constance 1949, 1950; Gillett 1968). As indicated m Fig. 1 and in 
the taxonomic treatment that follows, we recognize three infraspecific elements 

'Present address: U.S. Army Corps of Engineers, Planning Division, P.O. Box 1715, 
Baltimore, MD 21203-1715. 

in the group, all of which are distinctly allopatric and show little or no tendency 
to intergrade when observed in the field. The var. austrotexana, a segregate from 
yd.L patuliflora as treated by Constance, is probably closer to P. laxa Small than it 
is to P. patuliflora var. patuliflora [which appears equally close to P. strictiflora 
(Engelm. & Gray) A. Gray, according to Constance (1949)}- Indeed, Constance 
in his discussion of P. laxa, states that the species is: 

. . . exceedingly difficult to separate from P, patuliflora in the herbarium . . . since all the 
character of P. laxa, except perhaps the low number of ovules, have been diffused into P. 
patuliflora, its distinctness rests solely upon a combination of characters, none of which is 

He goes on to note, however, that in the field it is easily recognized by its small, 
pale flowers, petiolate cauline leaves, sparse pubescence and preference for shaded 
alluvial situations, all of which the senior author also observed in her field excur- 
sions. Nevertheless, Constance opined that P. laxa has been very nearly "swamped 
out" by the competition from its own recombinational products with P. patuliflora 
var. teucriifolia. Our field observations and examination of distributional ranges, 
make it difficult to accept that P. laxa has, to any considerable extent, hybrid- 
ized with P. patuliflora var. teucriifolia. Rather, the distribution pattern and mor- 
phological data strongly suggest that P. patuliflora var. austrotexana is a region- 
ally differentiated taxon little affected by hybridization from P. laxa or other 
taxa. Whatever its origin, it would appear on both morphological and geo- 
graphical grounds that P. laxa is much closer to P. patuliflora var. austrotexana 
than It is to P. patuliflora var. teucriifolia. 


The three varieties oiPhacelia patuliflora in Texas are distinctly separated geo- 
graphically (Fig. 1). Interestingly, the collective distribution of the subgenus 
Cosmanthus indicates a migration northward from Mexico (Constance 1950; 
Turner 1991). Within Texas this migration presumably allowed for geographi- 
cal diversification and habitat specialization for each variety. 

Phacelia patuliflora is characterized by hard-to-locate but often relatively dense 
populations which appear to be thriving. The species is not present in many 
apparently suitable areas near the known populations. Frequent human distur- 
bance or overgrazing at the sites where P. patuliflora does occur might explain 
the difficulty in locating sizeable colonies. 

Phacelia patuliflora (Engelm. & A. Gra' 
10:321. l?,15.Eucotapatuliflora'Enge\m. t 
Type: U.S. A: Texas: Austin Co.: in a woods 

OYER AND Turner, Systematics of Phacelia patuliflora 

Phacdui hispida Buckl., Proc. Acad. Nat. Sci. Philadelphia 13:463. 1862. nc 
1878. Type: U.S.A.: Texas: Travis Co.: Austin, I860, Buckley s.n. (holotype: P^ 
GH, MO). 

Peduncles with small, well-defined, glandular hairs, these ca 0.1 mm in length 
and dispersed among a rather even vestiture of mostly short, non-glandular 

2. Calyx lobes rounded or rather abuiptlv obtuse, mostly sandv soils of southern 

Texas var austrotexana 

Phacelia patuliflora (Engelm. & A. Gray) A. Gray var. patuliflora 

Annual herbs, 8-60 cm high, branching at the base with branches stiffly as- 
cending to decumbent. Leaves rarely rosulate or persistent, petiolate, truncate at 
base, oblong to oval, 2-10 cm long, 1-4 cm broad, pinnately lobed to pinnati- 
fid with 1-7 pairs of smaller lobes or remote leaflets at base and a larger terminal 
lobe, all coarsely dentate or lobed with obtuse or acute teeth, strigulose to hir- 
sute on both surfaces, the cauline leaves reduced, mainly sessile, oblong and 
coarsely dentate, distinctly lobed. Inflorescence a scorpoid cyme of 5-30 flowers, 
loosely hirsute to strigulose or canescent; the peduncles with small, well-de- 
fined glandular hairs, the latter about 0.1 mm in length; the mature pedicels 
weakly ascending to widely spreading or decumbent, 3-15 mm long. Corolla, 
purplish-violet to lavender, commonly with white centers, broadly campanu- 
late to rotate-campanulate, 8-16 mm broad, the lobes finely crenulate, moder- 
ately to sparsely pilose on back. Stamens 5-8 mm long, included, the anthers 
1 .0-1.5 mm long. Style included, cleft to midpoint, hirsute below. Capsule 
globose, 4-6 mm across, the seeds 10-15, about 2 mm in length, ovoid-angled, 
brown, areolate and finely alveolate. Chromosome number, n = 9 pairs. 

Distribution and Ecology: Central and eastern Texas (Fig. 1) mainly in sandy 
soils of roadsides, railroad embankments and alluvial soils of stream and river 
beds. The plants occur in disturbed areas in fuU sun or in partial shade of de- 
ciduous roadside thickets. Often grazed, plants begin to flower in mid-March, 
peak in early April and have set seed by the middle of May. 

Population Site Descriptions: {¥ig. 1) Three populations, 5-15 individuals each, 
of var. patuliflora were found in Burnet County on April 1 3 , 1 99 1 ■ Populations 
1 , 2 and 3 were located on sandy, well-drained roadside banks in the Buchanan 
Dam area. These sites were situated on large sandstone outcrops with smaller 
flags present on the surface. Some of the plants in all of the sites had been grazed. 







■'~ i 7 

1 i 

I pl T 1 



1 ,'"-'■' 



-'I A ■:--.- 



V\ " '' 

-;l\ "^ 

° var.teu 
^ var. pa 
• var. aus 



L^,e " 

n specimens at LL, TEX. Popula 

Interestingly, plants were confined to the sandy areas, although tracts with 
abundant humus were in close proximity. Grasses and occasional plants of 
Opuntia macrorhiza characterized the area. 

Population 4 of var. patuliflora was observed at or near the type locality 
in San Felipe, Texas on April 3, 1992. This population was rather dense, 
with 1000-1500 plants covering approximately half an acre. The plants 
were growing in very sandy loam with a few sandstone outcrops and were 
located in full sun. Other plants growing in this area were Lupinus texefuis, 
Pyrrhopappus pauaflorus, Gaillardia pulchella, Cinium texanum and Castilleja 

Mover and Turner, Systematics of Phaceliapatuliflora 249 

Phacelia patuliflora var. teucriifolia (I.M. Johnst.) Constance. Type: 
MEXICO. Coahuila: Muzquiz, 12 Apr 1936, Manh 2120 (holotype: GH; isotype: 
Annual herbs, 10-60 cm high, branching at the base with branches stiffly 
ascending. Leaves rarely rosulate or persistent, petiolate, truncate at base, ob- 
long to oval, 2-10 cm long, 1-4 cm broad, pinnately lobed to pinnatifid with 
1-7 pairs of smaller lobes or remote leaflets at base and a large terminal lobe, all 
coarsely dentate or lobed with obtuse or acute teeth, strigulose to hirsute on 
both surfaces, the cauline leaves reduced, mainly sessile, oblong and coarsely 
dentate, distinctly lobed. Inflorescence a scorpoid cyme of 5-30 flowers, loosely 
hirsute to strigulose or canescent, the mature pedicels weakly to stiffly ascend- 
ing, 3-1 5 mm long. Corolla, purplish-violet to deep bluish-purple, commonly 
with white centers, broadly campanulate to rotate-campanulate, 12-20 mm 
broad, the lobes finely crenulate, moderately to sparsely pilose on back. Stamens 
5-8 mm long, included, the anthers 1.0-1.5 mm long. Style included, cleft to 
midpoint, hirsute below. Capsule globose, 4-6 mm across, the seeds 10-15, 
about 2 mm in length, ovoid-angled, brown, areolate and finely alveolate. Chro- 

Dhtrihution and Ecology: Edwards Plateau region of Texas southward to Mexico 
(Figure 1). Plants commonly occur in calcareous soils in draws and river beds as 
well as along roadsides. Populations are frequently located in the partial shade 
of deciduous trees and in the full sun of disturbed roadside ditches. Flowering 
begins in late March with its peak in mid-April. Seeds are set and plants die 
back by late May. 

Population Site Descriptions: (Fig. 1) Populations 5 and 6 occurred in Montell, 
Uvalde County and were sampled in mid-April of 1 99 1 . Both were located on 
highway embankments. Populations were comprised of 20-50 plants which 
were widely scattered, located in ftill sun and on the naturally calcareous soil of 
the western Edwards Plateau region. Associated species included. Delphinium 
carolinianum, Callirhoe involucrata, Verbena bipinnatifida and Phlox drummondii, as 
well as assorted grasses. 

Populations 7, 8 and 9 were all located along the San Saba River in Menard 
County. These sites were characterized by calcareous soils of a grainy nature that 
seemed well drained. Populations 7, located at apicnic area on highway 83, two 
miles south of Menard, and population 8, located in the town of Menard, were 
situated in the partial shade of pecan bottoms. Both populations were dense, 
comprised of 500-1500 individuals. Other species in these areas included 
Oenothera speciosa, Gaura coccinea and Penstemon triflorus. 

Population 9 was located on the banks of a road cut along FM 1311 where it 
crosses the San Saba River and was comprised of 24 individuals. The soil at this 
site was sand that appeared to have been recently deposited. It was also in 

partial shade. Associated species were grasses and abtmdant individuals of 
Clematis pitcheri. 

Phacelia patuliflora var. austrotexana J.A. Moyer, var. nov. 

Differc a Phacelia patuliflora (Engelm. & A. Gray) A. Gray var. patuliflora trichomatibus 

glandulosis carentibus. 

Annual herbs, 8-60 cm high, branching at the base with branches stiffly 
ascending to decumbent. Leaves rarely rosulate or persistent, petiolate, truncate 
at base, oblong to oval, 2-10 cm long, 1-4 cm broad, pinnately lobed to pin- 
natifid with 1-7 pairs of smaller lobes or remote leaflets at base and a large 
terminal lobe, all coarsely dentate or lobed with obtuse or acute teeth, strigulose 
to hirsute on both surfaces, the cauline leaves reduced, mainly sessile, oblong 
and coarsely dentate, distinctly lobed. Inflorescence a scorpoid cyme of 5-30 
flowers, loosely hirsute to strigulose or canescent; the mature pedicels weakly 
ascending to widely spreading or decumbent, 3-15 mm long. Corolla, pur- 
plish-violet to lavender, commonly with white centers, broadly campanulate to 
rotate-campanulate, 7-1 6 mm broad, the lobes finely crenulate, moderately to 
sparsely pilose on back. Stamens 5-8 mm long, included, the anthers oblong 
1 .0-1.5 mm long. Style included, cleft to midpoint, hirsute below. Capsule 
globose, 4-6 mm across, the seeds 10-15, about 2 mm in length, ovoid-angled, 
brown, areolate and finely alveolate. Chromosome number, n=9 pairs. 

Type: U.S.A.: TEXAS: San Patricio Co.: along railroad ca. 2.5 mi. SW of Sinton, 22 Mar 
1969, D.S. Correll 36834 (holoi-ype: LL!; isotype: TEX!). 

Representative specimens: TEXAS. Bee Co.: just south of Pettus, 2 Mar 1968, Correll 333 1 1 . 
Brooks Co.: King Ranch, Encino Division, 26 Apr 1949, Lunckll 14975. Cameron Co.: Ar- 
royo Colorado, Harlingen, 1 3 Mar 1964, Correll 28973- Dimmit Co.: Carrizo Springs, 12 Mar 
1964, Cuellar43. Duval Co.: 7 mi W of Hebbronville, 5 Mar 1962, Tnvino4. Goliad Co.: 
open places, Mar 1927, C.B. Williams 45. Hidalgo Co.: S of Encino, 19 Mar 1942, Lundell 
10829. Jim Hogg Co.: Hebbronville, 14 Mar 193 1, Tharps.n. Jim Wells Co.: field at Alfred, 
30 Mar 1970, Correll 38310. Kenedy Co.: Sarita along Hwy 96, 14 Mar l9Al, Runyon 2469. 
Kleberg Co.: 10 mi S of KingsviUe, 6 Mar 1959, Turner 4467. Live Oak Co.: along US Hwy. 
59, 20 SW of Georgevi^est, 14 Mar 1964 Fowler & Vergara 99. Nueces Co.: Petronila Creek, 
between Bishop and Chapman Ranch, 12 Mar 1964, Correll 28925. San Patricio Co.: near 
Mathis, 29 Mar I94l, Lundell 10081. Webb Co.: 8.3 mi S of Mirando City, 16 Mar 1966, 
Correll 32259. 

Distribution and Ecology: South and southeastern Texas in the sandy, well- 
drained soils of roadsides and alluvial stream beds (Fig. 1). Found in the follow- 
ing counties: Bee, Brooks, Cameron, Dimmit, Duval, Goliad, Hidalgo, Jim 
Hogg, Jim Wells, Kenedy, Kleberg, Live Oak, Nueces, San Patricio, Webb. 
Populations were found most frequently in full sun and often comprised the 
only vegetation in highly disturbed areas. Flowering begins in March with 
maximum anthesis in early April. 

Population Site Descriptions: (Fig. 1) Population 10, located in Bee County, 

MoYHR AND Turner, Systematics of Phacelia patuliflora 251 

was sampled on 12 March 1991. It was comprised of approximately 500- 
700 plants which were found growing on the banks of a roadside drainage 
area in downtown Beeville. The drainage ditch walls and associated bank 
were comprised of sand with gravel and was located in the full sun. The 
population extended to a railroad embankment located approximately 20 
meters from the roadside area. This embankment, also comprised of sand 
and gravel, was located in full sun and was well-drained. Individuals were 
not found in the 20 meter swath of loamy sand that lay between the drain- 
age area and the embankment. Associated species found growing in the 
sand and gravel included Corydalis micrantha and isolated individuals of 
Opuntia macrorhiza. 

Population 1 1 was collected in Cameron County on 14 March 1991 • The site 
was in an abandoned commercial area off of the access road to Highway 77. 
Approximately 100-1 50 individuals were growing in what appeared to have 
been a sand and gravel parking lot in direct sun. No plants were found in a 
nearby (20 meters distance) overgrown garden area of rich loam. 

Kenedy County was visited on 14 March 1991 and three populations were 
located. Population 12 was located along Highway 77 on the sandy berm of a 
drainage area. The berm, located in full sun, appeared to have been deposited 
some years prior and was comprised entirely of sand. Approximately 50 indi- 
viduals were growing in association with Sisyrinchium ensigerum and varied grasses. 

Population 1 3 was the only partially cultivated population found. Several thou- 
sand individuals were growing in the sandy loam of Sari ta Square in the center 
of Sarita. These plants were being watered daily. All individuals were located in 
full sun in association with Gaillardia pulchella, Castilleja indivisa, Sisyrinchium 
ensigerum, Gaura coccinea, Argemone albiflora and varied herbs and grasses. 

Population 14 was located along a roadside and the railroad embankment 
running parallel to and approximately 15 meters from Highway 77 near Ytturia. 
This site was characterized by extremely dry, well-drained sand and gravel. 
Approximately 100-200 individuals were present with only sparse grasses. 

Three populations were found in Kleberg County on 14 March 1991 ■ Popu- 
lation 1 5 was comprised of 1 5 plants and was located on the alluvial sand of 
Escondido Creek just south of Kingsville. Individuals were found only in the 
sand of the banks, and not in the surrounding grassy field. All individuals were 
located in full sun. Associated species included Buchnerafloridana and Phlox 

Population 16 occurred on a railroad embankment along Highway 77 at its 
junction with FM 1 1 18 in alluvial sand and gravel. The 22 plants observed were 
in hall sun and were widely separated. Associated vegetation was comprised of 
Corydalis micrantha, Cirsium texanum, Opuntia macrorhiza, and Sisyrinchium ensigerum. 

Population 17 was also growing on a railroad embankment running parallel 
to Highway 77 just south of its junction with FM 628, in gravelly sand. 

252 SiDA 16(2) 1994 

This population was widely scattered and was comprised of approximately 
50-60 individuals. However, this group was in the partial shade of a large 
Acacia grove. Corydalis mtcrantha and Sisyrinchium ensigerum were the only 
other species found on the embankment. 

Population 18 was located in Nueces County and was sampled on 13 
March 1991. It was comprised of 25-50 individuals growing in sand at the 
foundation of a roadside warehouse off of the Padre Island Expressway in 
Flour Bluff. The individuals were restricted to the sandy soils that appeared 
to have been placed at the foundation for the purpose of creating a drainage 
bank. The site was located in full sun; grasses were the only other species 

The present study represents part of a Master's thesis by the senior author. It 
is based upon field work, as indicated, and examination of approximately 300 
specimens of the taxa concerned on file in the Plant Resources Center at the 
University of Texas at Austin (LL, TEX). We are grateful to Guy Nesom for the 
Latin diagnosis and to the McLean Family, Red Stoner and Marky Miles for 
their generous donation of time and effort in locating populations. 

ubgenus Cosmanthus (Hydrophyila( 

.. 1950. Some interspecific relationships in Phacelia subgenus Cosmanthus. Proc. Amer. 
rts 78:135-147. 

i.W. 1968. Systematic relationships in the Cosmanthus and Phacelias (Hydrophyl- 
Brittonia 20:368-374. 



Department of Botany and Range Science 
Brigham Young University 
Pravo, UT 84602, U.S.A. 

MHlerocaulis Erasmus ex Tidwell was established as a new genus for Miller's "Osmundacaulis 
herktii group. " Amtralosmunda Hill, Forsyth & Green was proposed subsequently for osmunda- 
ceous stems anatomically similar to MHlerocaulis, but without leaf gaps. However, a few species 

Amtralosmunda is placed in the synonymy oi MHlerocaulis. MHlerocaulis is emended to include 
those osmundaceous species without or having only an occasional leaf gap. Under the new 
arrangement, MHlerocaulis includes four species M. dunlopii, M. indica, M. indentata, and M. 
chubutensis. A new genus, Ashicaulis, is proposed for species formerly in MHlerocaidis having 
numerous leaf gaps. It contains 22 species, all of which are recognized nomenclaturally by 

MHlerocaulis Erasmus ex Tidwell se establecio como un nuevo genero para el grupo de Miller 
')smundacatdis herbstii" . Se propuso por ello Amtralosmunda Hill, Forsyth & Green para los tallos 
;mundaceos anatomicamente similares a MHlerocaulis, pero sin lagunas foliates. Sin embargo, 
iluyendo su tipo, 7W. £^a«/o/'iz, carecen de o tienen ocasional- 

The genus MHlerocaulis was established as a new taxon based upon the infor- 
mal "Osmundacaulis herbstii group" that was originally discussed by Miller (1967). 
The ^^Osmundacaulis herbstii group" was inefifectively published (Art. 29) as a new 
genus MHlerocaulis by Erasmus in his unpublished doctoral dissertation (1978). 
Herbst (1 98 1:37) accepted Erasmus' MUlerocaults but failed to validate it with a 
description (Art. 32.1)" (Tidwell 1986:402). MHlerocaulis was validated by 
Tidwell (1986). 

Australosmunda was proposed for osmundaceous stems similar to MHlerocaidis, 
but without leaf gaps (Hill et al. 1989; Fig. 1 A). However, there are afew species 
in MHlerocaulis, including its type, Al. dunlopii (Kidston and Gwynne-Vaughan 

^^ '«, « 

K ^'S 


linders of four species oi Milleroca. 
ndentata {Australosmunda indentata), 
ni B, Part of continuous xylem cyl 

lack of leaf gaps. B, C, D. Millerocaulis dunlopti B, Part of continuous xylem cylinder, C, D, 

ing many leaf gaps in their xylem cylinders. G. Millerocaulis indica with continuous xylem 
cylinder and forming leaf trace. H, I, J, K, Millerocaulis chubutmsis, H. xylem cylinder without 
leaf gaps, I, J, K, leaf trace formation and departure without leavmg gaps. (Bars = 1mm). (A. 
Drawn from holotype oi Austalosmunda (=M.) indentata. Hill et al. 1989; B, C, D, Drawn from 
holotype of M. dunlopti, Kidston & Gwynne-Vaughan 1907; E. Redrawn from Archangelsky & 
de la Sota 1963; E Drawn from holotype oi Ashicaults wadet, Tidwell & Rushforth 1968; G. 
Redrawn from Sharma 1973; H. Drawn from holotype of M. chubutensis, Herbst 1977, 1,J,K. 
Redrawn from Herbst 1977). 

Tjdwell, Ashicaulis, a new genus in the Osmundaceae 255 

1907; Figs. IB, C, D), that also lack or have an occasional leaf gap. Since the 
type oi Milkrocaidis essentially lacks leaf gaps, Australosmunda is placed in the 
synonymy oi Millerocaulh , and Millerocaulis is restricted to fossil osmundaceous 
axes lacking or having very rare or only an occasional ill-deflned leaf gap. Ashi- 
caulis is proposed for those species formerly in Millerocaulis that have numerous 
deflmteieafgaps(Figs. 1E,F). 

Millerocaulis Erasmus ex Tidwell emend. Tidwell 

Fossil osmundaceous rhizomes or arborescent axes containing stem or branch- 
ing stems surrounded by a mantle of leaf bases and roots. Stele an ectophloic 
siphonostele, with xylem cylinder usually continuous, generally 3-1 5 , some- 
times up to 20, tracheids thick radially, with leaf gaps either totally lacking, 
incomplete, or very rare. Incomplete leaf gaps extending up to three-quarters 
through the metaxylem cylinder. Leaf trace separating from the xylem cylinder 
with only one protoxylem cluster and often, but not always, lacking axillary 
sclerenchyma. Scelerotic outer cortex of stem usually much wider than paren- 
chymatous inner cortex. Petiole bases stipulate, and adventitious roots arising 
either singly or in pairs. 

Type: M. dunlopii (Kidston & Gwynne-Vaughan) Tidwell (Basionym: Osmundttes dunlopii 

Millerocaulis chubutensis (Herbst) Tidwell, comb. nov. Basionym: OsmimdacauUs 

Millerocaulis dunlopii (Kidston & Gwynne-Vaughan) Tidwell, comb. nov. 

Basionym: Osmundites dunlopii Kidston & Gwynne-Vaughan, Trans. Roy. Soc. Edinburgh 

45:759. 1907 {"dunlopi"). Osmundacaulis dunlopii (Kidston & Gwynne-Vaughan) Miller, 

Contr. Mus. Paleo. Univ. Michigan [21:146. 1967 i:'dunlopi"), nom. invalid, under Art. 

33.2 — no page reference to basionym} 23:135. 1971. Osmundites aucklandicus Marshall, 

Trans. & Proc. New Zealand Inst. 56:210. 1924. 
Millerocaulis indentata (Hill, Forsyth & Green) Tidwell, comb. nov. Basionym: 

Australosmunda indentata Hill, Forsyth & Green, Palaeontology 32:292. 1989. 
Millerocaulis indica (Sharma) Tidwell, comb. nov. Basionym: Osmundacaulis indica 

Sharma, Palaeontographica l40B: 157. 1973. 

Ashicaulis Tidwell, gen. nov. 

Fossil osmundaceous rhizomes, rarely arborescent axes, containing stem or 
branching stems surrounded by a mantle of leaf bases and roots. Stele an ec- 
tophloic-dictyoxylic-siphonostele (Miller 197 1), with a xylem cylinder up to 
20 tracheids thick, with many definite leaf gaps. Leaf trace separating from the 
xylem cylinder with only one protoxylem cluster and often, but not always, 
lacking axillary sclerenchyma. Commonly, the outer sclerenchymatous cortex is 
wider than the parenchymatous inner, rarely the same width. Petiole bases stipular 
and adventitious roots arising either singly or in pairs. 

TYPE: Ashkaulis herhsth (Archangelsky & de la Sota) Tidwcll {Osmundites herbsth Archangel- 

The generic name honors my good friend and colleague Professor Sidney R. 
Ash of Weber State University of Ogden, Utah, in recognition of his numerous 
significant contributions to paleobotany. The list of species assigned to Ashkaulis 
and their synonyms are as follows: 
Ashicaulis amajolensis (Sharma) Tidwell, comb. nov. Basionym: Osmundacaulh 

amajolensisSh^tma., Palaeontographica HOB: 156. 1973. MilkrocauUs amajokmis {ShdiTma) 

Tidwell,Sida 11:402. 1987. 
Ashkaulis beardmorensis (SchopO Tidwell, comb. nov. Basionym: Ostnundacaulis 

beardmarensisSchopiGin.J. Bot. 56:3034. 1978. MiilerocauHs beardmorensis (Schopf)Tidwel\, 

Ashicaulis broganii (Tidwell, Munzing & Banks) Tidwell, comb. nov. Basionym: 
Millerocaulis broganii Tidwell, Munzing & Banks, Palaeontographica 223B:98. 1991 . 

Ashicaulis estipularis (Sharma, Bohr & Singh) Tidwell, comb. nov. Ba.sionym: 
Osmundacaulis estipularis Sharma, Bohra & Singh, Phytomorphology 8:61 . 1979 ("estipu- 
lare"). Millerocaulis estipularis (Sharma, Bohra & Singh) Tidwell, Sida 1 1 :403. 1987. 

Ashicaulis gibbiana (Kidston & Gwynne-Vaughan) Tidwell, comb. nov. 
Basionym: Osmundites gibbiana Kidston & Gwynne-Vaughan, Trans. Roy. Soc. Edmburgh 
45:763. 1907. Osmundacaulis gibbiana (Kidston & Gwynne-Vaughan) Miller, Concr. Mus. 

reference to basionym} 23: 1 36. 197 1 . Millerocaulis gibbiana (Kidston & Gwynne-Vaughan) 

Tidwell, Sida 11:403. 1987. 
Ashicaulis guptai (Sharma) Tidwell, comb. nov. Basionym: Osmundacaulis guptai 

Sharma, Palaeontographica 140B: 1 54. 1973. Millerocaulis guptai (Sharma) Tidwell, Sida 

11:403. 1987. 
Ashicaulis hebeiensis (Wang) Tidwell, comb. nov. Basionym: Osmundacaulis 

hebetensis Wang, Rev. Palaeobot. Palynol. 39:93. 1983. Millerocaulis hebeiensis (Wang) 

Tidwell,Sida 11:403. 1987. 
Ashicaulis herbstii (Archangelsky & de la Sota) Tidwell, comb. nov. Basionym: 

Osmundites herbstii Archangelsky & de la Sota, Ameghiniana 3:135.1963- Osmundacaulis 

herbstti (Archangelsky & de la Sota) Miller, Contr. Mus. Paleontol. Univ. Michigan (2 1 : 146. 

1967,nom.mvalid. under Art. 33.2 — no page reference to basionym) 23: 134. 1971. 

Millerocaulis herbstii (Archangelsky & de la Sota) Tidwell, Sida 1 1 :403. 1987. 
Ashicaulis johnstonii (Tidwell, Munzing & Banks) Tidwell, comb. nov. 

Basionym: MilleivcaulisjohnstoniiTid^dl Munzmg & Banks, Palaeontographica 223B:94. 

Ashicaulis kidstonii (Stopes) Tidwell, comb. nov. Basionym: Osmundites kidstonii 

Stopes, Ann. Bot. 35:55. 1921 {"kidstom"). Osmundacaulis kidstonii {Stopfts)M\\\fiv, Contr. 

Mus. Paleontol. Univ. Michigan (21:146. 1967 {"kidstoni"), nom. invalid, under Art. 33.2 

Ashicaulis liaoningensis (Wu & Shao-lin) Tidwell, 

Millerocaulis liaoningensis Wu & Shao-lin, Acta Palaeontol. Sine 

TiDWELL, Ashicaulis, a new genus in the Osmundaceae 257 

Ashicaulis patagonica (Archangelsky & de la Sota) Tidwell, comb. nov. 

Basionym; Osmundites patagonica Archangelsky & de la Sota, Ameghiniana 2:153. 1962. 
Osmundacaulis patagomca (Archangelsky & de la Sota) Miller, Contr. Mus. Paleotol. Univ. 
Michigan [21:146. 1967, nom. invalid, under Art. 33.2 — no page reference to ba- 
sionym] 23: 1 36. 1 97 1 . Millerocaulis patagomca (Archangelsky & de la Sota) Tidwell, Sida 
Ashicaulis rajmahalensis (Gupta) Tidwell, comb. nov. Basionym: Osmundites 
rajmahalensis GuY>t2i, Proc. Ind. Sci. Congr. Varanasi 55:428. 1968. Osmundites rajmahalen- 
sis Gupta, Palaeontographica 1 30B: 1 74. 1 970. Osmundacaulis rajmahalensis (Gupta) Sharma, 
Palaeontographica l40B:152. 1973. Millerocaulis rajmahalensis (Gupta) Tidwell, Sida 

Ashicaulis sahnii (Mittre) Tidwell, 

Paleobotanist4:113. 1955. Osmundacaulis sahnii (M:^x.nt)Mv\\^T, Contr. Mus. PalaeontoL 
Univ. Michigan [21:146. 1967, nom. invalid, under Art. 33.2 — no page reference to 
basionym} 23: 1 3 5 . 1 97 1 . Millerocaulis sahnii (Mittre) Tidwell, Sida 1 1 :403 . 1 987. 

Ashicaulis santaecrusis (Herbst) Tidwell, comb. nov. Basionym: Osmundacaulis 
santaecrusis Herbst, Facena 1:21.1977. 

Ashicaulis spinksii (Tidwell, Munzing & Banks) Tidwell, comb. nov. Ba- 
sionym: Millerocaulis spinksii Tidwell, Munzing & Banks, Palaeontographica 223B:96. 

Ashicaulis swanensis (Tidwell, Munzing & Banks) Tidwell, comb. nov. 
Basionym: Millerocaulis swanensisTidv^eW, Munzing & Banks, Palaeontographica 223B:99. 

Ashicauhs wadei (Tidwell & Rushforth) Tidwell, comb. nov. Basionym: Oj- 
mundacaulis ivadeiTidv/tW & Rushforth, Bull. Torrey Bot. Club 97: 137. 1970. Millerocaulis 
ivadet (Tidwell & Rushforth), Tidwell, Sida 1 1 :403. 1987. 

Ashicaulis websterii (Tidwell, Munzing & Banks) Tidwell, comb. nov. Ba- 
sionym: Millerocaulis websterii Tidwell, Munzing & Banks, Palaeontographica 223B: 97. 

Hill et al. ( 1 989) considered Australosnmnda tndentata the first fossil osmun- 
daceous rhizome described with a simple siphonostele and a parenchymatous 
pith without realizing that the holotype oi Millerocaulis , M. dunlopii, also has a 
simple siphonostele. The reason is the considerable confusion surrounding the 
presence or absence of leaf gaps in M. dunlopii. As pointed out by Kidston and 
Gwynne-Vaughan (1907) in describing the holotype, "The most important 
anatomical character of this species is the almost complete absence of leaf gaps 
in the xylem ring of the stem. The fact that most of the leaf traces, if not all, 
depart without in any way interrupting the continuity of the xylem ring, so that 
the "medullary rays" characteristic of the osmundaceous stele in general are 
almost or completely absent. Most of these breaks in the xylem ring are clearly 

due to accident, but it is just possible that some of them may liave been occu- 
pied by thin-walled cells which decayed before fossilization. Even if such med- 
ullary rays actually were present in the living plant, they must have been ex- 
tremely narrow and very rare." "... and reference to the longitudinal sections 
will make it clear that no medullary ray is caused by the departure of the xylem 
of the leaf-trace and that the continuity of the deeper portion of the xylem ring 
is undisturbed." (p. 760-761). Thus, this species lacks leaf gaps. 

Sinnott (1910), however, attributed the apparent lack of leaf gaps in the 
holotype of M. dunlopii to its having very short and narrow gaps that were oblit- 
erated by the process of fossilization, an intriguing explanation but without any 
real evidence. It is interesting to note that Sinnott also illustrated a diverging 
leaf trace for Osmunda regain in an "apparently gapless fashion" and one for 0. 
cinnamomea and others for Todea superba and X hymenophyllokks that had not formed 
complete gaps. The lack of leaf gaps in extant Osmundaceae is rare. Miller (pers. 
comm. 1993) mentioned that stems of modern osmundaceous taxa may be with- 
out complete leaf gaps near branches. 

Several osmundaceous specimens with leaf gaps have been reported as Osmun- 
dttes{= Millerocaulis) dimlopii from the Jurassic of New Zealand (Sinnott 1914) 
and the Jurassic near Purga, Queensland (Sahni 1920). However, none appear to 
be that species. One of Sinnott 's specimens, for example, is most likely Ashkau- 
Ih wrightti described from Tasmania (Tidwell et al. 1991). His other specimens 
also belong to Ashtcaulis but are too poorly preserved to be identifiable to spe- 
cies. Of the five leaf gaps Sahni (1920) noted in the transverse section of the 
stem of one of the four incomplete specimens he attributed to 0.(=M.) dunlopii, 
two very narrow gaps may be valid, but even these, could be due to fracturing 
prior to preservation. None of his alleged gaps have traces opposite them. Sahni 
further stated, "In the only longitudinal section that was made, three leaf-traces 
are seen arising, but while it is impossible to say whether all of these have been 
cut medially, in no case is the cauline strand interrupted opposite an off-coming 
leaf trace" (p. 1 3). Furthermore, it is doubtful if the specimen (#140) Sahni (1 920) 
illustrated as text - fig. 2 is M. dunlopii, but more likely represents a species of 
Ashicaulis. He founded much of his identification on the curved leaf traces, scle- 
rotic rings, and "the two large sclerotic strands inside the curved ends of the 
leaf-trace, which is characteristic of 0. dunlopit." He used these criteria based on 
the limited knowledge of the Osmundaceae in Australia at the time. It is now 
known that the first two characters occur in essentially all the extant and extinct 
species of the Osmundaceae (Hewitson 1962; Miller 1 967 , 1 97 1 ) and the last 
feature in nearly all described species oi Millerocaulis and Ashicaulis from that 
continent (Tidwell et al. 1991). 

The distribution of sclerenchyma in the stipular wings, which has proven to 
be useful in species determination (Hewitson 1962; Miller 1967), was largely 
ignored in the specimens reported by Sahni (1920) and Sinnott (1910, 1914) as 

TiDWELL, Ashicaulis , a new genus in the Osmundaceae 259 

Osmundite5{=M.) dunlopii. The distribution in the wings of one petiole figured 
by Sahni (pi. I, fig.4) was noted to be like that described for M. dunlopii, but that 
particular petiole was not attached to any stem. The stem Sahni ( 1 920) figured 
(pi. I, fig. 3) had attached petiole bases, but none was figured or described. Con- 
sequently, it is doubtful whether any were really M. dunlopii. 

Edwards (1933) pointed out in discussing his fossil osmundaceous specimen 
from central Australia that it had a continuous or almost continuous xylem 
cylinder that was not broken into separate strands similar to the Jurassic species 
oWsmundites {=Millerocaulis) dunlopii. Medwell (1954), Sharma (1973), and 
Andrews ( 1 961) all stated or implied that 0. (=7VI.) dunlopii possessed a continu- 
ous xylem ring, meaning no leaf gaps. 

Miller (1967, 1971) in his papers on the Osmundaceae z-cce^ittd the fossils of 
Smnott (1910, 1914) and Sahni (1920) as specimens of 0. (=M.) ^«^%V and 
concluded that the species had very narrow, some immediate, some delayed and 
some incomplete leaf gaps. In addition, he mentioned that the near absence of 
gaps in the type specimen of this species is not typical for the species. More 
recently, however. Miller (pers. comm. 1993) commented that in reviewing his 
notes he had not observed any evidence of leaf gaps when examining slides 
(# 1 242-1247) of the holotype of M . dunlopii from Gore (Otago), New Zealand 
at the British Museum of Natural History. He further stated that the holotype 
of M. dunlopii does not have leaf gaps, and that there was never any indication of 
the short narrow leaf gaps that might have been obliterated in the process of 
fossilization as per Sinnott's (1910) explanation. Furthermore, Miller noted that 
Kidston and Gwynne-Vaughan's (1907) photos (pi. I, fig. 3; pi. II, figs. 4-5) do 
not show any 'obliteration' of the xylem tracheids either. In conclusion, the only 
definite specimen of M. dunlopii known, is its holotype, and it lacks leaf gaps. 
Consequently, Australosmunda indmtata is reassigned to Millerocaulis . Millerocaulis 
dunlopii, the type, and M. indica, which also lacks leaf gaps (Sharma 1973; Fig. 
IG), are retained in this genus. Based on photographs oWsmundacaulis chubuten- 
sis from Argentina (Herbst 1977), this species also appears to be without leaf 
gaps (Figs. 1 H, I, J, K) and is placed in Millerocaulis as well. 

Millerocaulis, as now defined, with a Triassic species {M. indentata) and three 
Jurassic species illustrates another phylogenetic line in the Osmundaceae that is 
presently known only from the Southern Hemisphere. Osmundaceous taxa of 
this phylogenetic line in lacking gaps and having parenchymatous piths lend 
support to the hypothesis that evolution of the parenchymatous pith and evolu- 
tion of the leaf gap in the xylem cylinder were independent (Hill et al. 1989). 

The other species previously assigned to Millerocaulis have numerous definite 
leaf gaps are hereby transferred to the new genus Ashicaulis. Ashicaulis, presently 
composed of 22 species, is known from both the Northern and Southern Hemi- 
spheres and is currently known from the Triassic to Early Cretaceous. 

I am grateful to Dr. Dan H. Nicolson of the Smithsonian Institution, Dr. 
Charles N. Miller, Jr. of the University of Montana, Dr. Alfred Traverse of Penn 
State University, and Dr. Sidney R. Ash of Weber State University for their 
assistance with nomenclatural problems as well as reviewing earlier versions of 
this paper. 


Archangelsky, S. and E.R. De La Soto. 1962. Estudio anatomico de un estipite petrificado do 
"Osmundites," de edad Jurasica, procedente del Gran Bajo de San Julian, Provincia de Santa 
Cruz. Ameghiniana 2 : 1 5 3- 1 64 . 

1963. Osmundites herbstii, nueve petrificacion Triasica de el Tranquilo, Provincia de 

Santa Cruz. Ameghiniana 3:135-140. 

Edwards, W.N. 1933. Osmundites from central Australia. Ann. Mag. Nat. His. 1 1:661-663. 

Erasmus, T. 1978. The anatomy and evolution oiOsmundacaulis Miller emend, with notes on 

unpublished Ph.D. dissertation. 

!. On a new species 

Gupta, K.M. 1970. Investi 



o{ Osmundites, 0. rajmak 

./.«.. sp.nov. from, 

:he Rajmahal Hills, Bih 

1306: 173-180. 

Herbst,R. 1977. Dos nuev 

as especies de Osmui 


de Osmundales de Arger 

itina. Facenal:19^ 


Herbst,R. 1981.G//tf/r&7/« 



{sensulato) del Permico su 

iperior de Paraguay 

Ameghiniana 18:35-50 

Hewitson,W.H. 1962.Cc 

)mparative morphology of the Osmundacea 

Gard. 49:59-93. 

ndF Green 1989 A newgenus of osmunc 

Upper Triassic of Tasmai 

iia. Palaeontology 3 



07. On the fossil Osmui 

Roy. Soc. Edinburgh 45: 


Marshall, R 1924. A new 

species oiOsmundttes from Kawhai, New Z< 

New Zealand Inst. 56:21 


Medwell,L.M. 1954. Are' 

.lew and revision of 

the flora of the Victoriar 

Roy. Soc. Victoria 65 :63- 


Miller, C.N. 1967. Evoluti 

on of the fern genus 

Osmunda. Contr. Mus. Vi 

gan 21:139-203. 

Miller, C.N. 1971. Evolut 

Univ. Michigan 23: 


Mittre,V. 1955.0jOT«Wi/e 

y sahnii sp. nov., a new species of petrified os. 

from India. Palaeobotani 

St 4: 113-118. 

Sahni, B. 1920. Petrified plant remains from i 

:he Queensland Mesozo 

TiDWELL, Ashicaulis, a new genus in the Osmundaceae 

, D. BoHRA, and R. Singh. 1979. Osmundacaulis estipulare sp. nov., a pet 

zome ffom Jurassic of Rajmahal Hills, India. Phytomorphology 29:46-52. 
SiNNOTT, E.W. 1910. Foliar gaps in the Osmundaceae. Ann. Bot. (Oxford) 24:107- 
Stopes, M.C. 1921 . The missing link in Osmundttes. Ann. Bot. (Oxford) 35:55-61. 
TiDWELL, W.D. 1986. MiUerocaulis, a new genus with species formerly in Osmundaca. 

(fossils: Osmundaceae). Sida 1 1:401^05. 
TiDWELL, W.D. 1992. MiUerocaulis richmondti sp. nov., an osmundaceous fern from 

strata near Little Swanport, Tasmania. Pap. & Proc. Roy. Soc. Tasmania 126:1-7. 
and S.R. RUSHFORTH. 1970. Osmundacaulis wadei, a new osmundaceous sp 

the Morrison Formation Qurassic) of Utah. Bull. Torrey Bot. Club 97:1 37-144. 
, G.E. MuNZiNG and M.R. Banks. 1991 . MiUerocaulis species (Osmundac 

Tasmania, Australia. Palaeontographica 223B:91-105. 
Wang, Z. 1983. Osmundacaulis hebeiensis, a new species of fossil rhizomes from tt 

Jurassic of China. Rev. Palaeobot. and Palynol. 39:87-107. 
Wu, Z. and Z. Shao-Lin. 1991 . A new species of osmundaceous rhizome from Midd 

of Liaoning, China. Acta Palaeontol. Sin. 3:7 14-727. 


^VERITT, James H. and D. Lynn Drawe. 1993. Trees, Shrubs & Cacti of 
South Texas. Texas Tech University Press, Lubbock, TX 79409-1037 
(1-800-832-4042). ISBN 0-89672-252X; ISBN 0-89672-253-8 
(pbk). $18.95 pbk. Text of colored photographs. 

lajority of trees, shrubs, and cacti occurring in clie 14 southernmost counties of Texas." 

;nough, mostly two per page to provide 
1 browsing through the booi<: ("Bairharis," 

Burt, Page. 1991. Barrenland Beauties. (ISBN 0-919315-25-9, pbk). 
Outcrop Ltd., The Northern Publishers, Box 1350, Yellowknife, 
N.W.T., Canada XIA 2N9. $25.00 (Canadian; includes postage). 

Dhotos — about 200 of them, a few full-page but most grouped two to four per page — ; 
excellent. Notes on habitat, ecology, and uses are included (with summaries in Inuinaktun, t 
written form of the dialect commonly spoken in the central arctic). Described, too, are t 

Jibliography, and an index to common and scientific names. Page Burt is a naturalist at Bathu 

nend the lodge highly). Although her fine book was written to fill the need for a guide to t 
narvelous, low-arctic flora of the inlet, its usefulness extends from there to Ellesmere Islar 



Botanical Research Institute of Texas 

309 Pecan Street 

Fort Worth, TX 76102-4060, U.S.A. 

Jew taxa of paleocropical Myrsinaceae are described from the Philippines and New Guinea 

species of Theaceae, probably Adinandra, which is not a myrsinaceous plant. The subge- 
c position oi Ardisia mystica is established. Ardtsia brassii Stone (non Sleumer) is renamed 

ueva Guinea. Se descubre que Ardisic 
a Theaceae, posiblemente un miembn 

This paper is a continuation of a series of studies of paieotropical Myrsinaceae 
from the Malesian region. Earlier papers in the series are included in the refer- 
ences. This paper is also Contribution no. 2 in the Precursors series for the Flora 
of the Philippines project. 

In the course of revising the Myrsinaceae in the Malesian region and in col- 
lecting intensively in the Philippines under the Philippine Plant Inventory project 
(PPI), new taxa, new biological and distributional data necessitate taxonomic 
and nomenclatural changes. New taxa are described mArdisia, Fittingia, and 
Tapeinosperma. Also, one purported variety of ArJisia is in fact theaceous and 
some further notes on species ofArdisia are included. 

A new species of Ardisia Swartz 

Ardisia reynosoi B.C. Stone, sp. nov. (Fig. 1). 

1:263-272. 1994 

rdisia reynosoi Sto 

ne (drawn from holotype). A. Leafy flowermg branchlet. B. Single 

Detached calyx 

2 lobes cut ofD, adaxial view. D. Corolla lobe, interior view; glands 

3ex, papillae abov 

e small staminal attachment. E. Stamen, abaxial view, with glandu- 

ctive.F. Ovary an 

d style. G. Placenta, the lower figure much enlarged to show rows of 

Trichomes from 

alyx margin. 

;, glabris, alat 

■evolutis; costa infra elevato; venis 
nifestis vel subobscuris; glandulis 
;, 2-3-floriferis, pedunculo 9-1 1 

usque ad 4 mm longis, glabris, g 
atis adnatis compressibus); stamii 
ctivo dorso bilinealiter glandu 
o 1 .5-2(^) mm longo producto 

Mt. Giting-giting, ridge to iVfa; 

.ince): Magdiwang, weste 
type: PNH; isotype: E). 

Paratypes: SIBUYAN: ridge to Mayos Peak, 1300 rr 
it, 24 Aug 1989, Argent & Reynoso 89-87 (E, PNH); ; 
t., dwarf shrub, 5 Jun 1992, B.C. Stone & E. Reynoso PPI 6923 (L, PNH, US). 

, submontane foi 
^ locality, 1400 r 

Stone, MalesianMyrsinaceae 265 

Etymology: Named for the collector Ernesto Reynoso whose extensive experi- 
ence in the Philippines qualified him as a team leader in the Philippine Plant 
Inventory Project. Many botanists and other collectors have been guided by 
"Estong" over the years, so this dedication is well deserved. 

Notes: This small, shrubby Ardisia occurs in elfin fi^rest and partly open areas 
on the steep upper ridges of Mt. Giting-giting; so far it is only known from 
there and is apparently endemic. The side-branches (usually 1 5-20 cm long) are 
ascending and bear small crowded ascending leaves. These may reach 8 cm but 
are usually shorter; the midrib beneath is tawny brown. The inflorescences are 
axillary and terminal, umbelliform, with umbels usually few (2-3) flowered; in 
fruit the calyx lobes attain a length of almost 2 mm and are slightly imbricate at 
base. Ripe fruits are dark purple, conspicuously obloid, and are about 5 mm 
long and 7 mm wide. In general appearance this species is very similar to A. 
gitingenm Elmer. 

The infrageneric position oi Ardisia mystica 

Ardisia mystica Stone, Malayan Forester 45: 114, fig. 5. 1982. 

This species was described from imperfect material and there was doubt as to 
which of the subgenera it pertains to. After examining type material o( Ardisia 
chrysophyllifolia King & Gamble (K) it was evident that the two species were 
closely related and that A. mystica had to belong in the same subgenus. Both in 
fact belong to Subg. Pimelandra. 

The ovary oi Ardisia mystica is minutely tomentellous and the flowering pedicels 
are about 1 mm long; while in A. chrysophyllifolia the ovary is glabrous and the 
flowering pedicels are 5 mm long. 

The identity oi Ardisia anceps var. borneensis Scheff. 

Ardisia anceps var. bomeenis Schefif., Myrs. Archip. Ind. 79- 1865 . 'Tolia sessilia 
denticulata longiuscule abrupte acimiinata, in sicco rubescentia." (Fig. 2). 

The type specimen bearing this name {Korthals s.n. from Gunong Rantau, 
South Borneo, in L!) is not myrsinaceous, but appears to be theaceous, probably 
a species oiAdinandra. It has acicular pellucid stramineous unicellular trichomes; 
the crenate-denticulate leaf margins with the distinct teeth are extensions of the 
veinlets. The habit of the leafy branchlets somewhat resembles an Ardisia with 
slight ridges (alulae) as prolongations of the petiole bases, and a pattern of glands 
on the leaf lamina. In several herbria, specimens oiAdinandra and other similar 
genera of Theaceae have been found lodged among undetermined material of 
Myrsinaceae, showing that there is a possibility of confusion among specimens 
of these families. The trichomes of Myrsinaceae are often ferrugineous or 
brown, seldom pale tawny, and the species with denticulate leaves have 

Stone, Malesian Myrsinaceae 267 

more delicate teeth with a different relationship to the veins. Scheffer him- 
self was doubtful as to the affinity of this specimen, stating "An Ardiual 
Nee fructiferam nee floriferam vidi. — Rami ancipites sunt. — Mynima ex 

A new name for Ardhta brass i/ Stone 

Ardisia brassiella B.C. Stone, nom. nov. A. brasm Stone, Proc. Natl. Acad. Sci. U.S.A. 

142: 52. 1990 (nonSIeumer.Blumea 33:130. 1988). Type: PAPUA NEW GUINEA: 

Sudest Island; Mt. Riu, W slopes, 300 m, 31 Aug 1956 (fl, fr), Brass 27934 (holotype: 


The name A. brassii Stone was inadvertently used in my 1990 study for the 

species represenred by Brass 27934 and 27825 , an error for A, brassiella which 

was intended, as Sleumer's earlier use of A, brassii (for a New Guinea species) 

preempts that epithet. 

Fittingia Mez 

This genus is endemic to New Guinea. The following new species is the sixth 
known; Sleumer (1988) recognized five species. 
Fittingia mariae B.C. Stone, sp. nov. (Figs. 3, 4). 

iatis, novellis subglabris vel sparsiter lepidotis. Folia anguste elliptico-ianceolatae, usque ad 
cm longae, 4 cm latae, integerrimae, chartacea, utt inque sensim acuminato-acutatae, infra 

submaniresto. Intlorescentiae in axillas toliorum ramulorum, breviter angustequ 
paucipaniculatae ad 5-6 cm longae, axibus omnibus pergracilibus et sparsiter lepidotulis 
pedunculis brevibus singularibus vel paucifasciculatis, ramis ad 10-12 mm longis, suboppositis 
bracteis subulatis glandulosis 1-2 mm longis, pedicellis ca. 2 mm longis, cymis l-3-floriferi5 
floribus ca. 2 mm longis, 4-meris, probabiliter omnino unisexualibus (masculis solis hi 
descriptis), calyce glabro cupulato 1.3 mm longo, lobis ovatis mtegerrimis multiglandulosi 

(0.5 mm) coalitis, extus conspicue atroglandulosis, intus omnino dense papulosis; filament! 
subteretibus ca. 0.5 mm longis, e basi lobi coroUae productis; antheris breviter oblongis fere 
mm longis apice emarginatis, inornatis, pallidis, eglandulosis; gynoecio rudimentale stylo 1. 

Type: NEW GUINEA. Morobe Province: Boana, 1,219 m in mossy montane scrub, 
slender shrublet, flowers dull pinkish, 4 Jul 1940, M.S. Clemens 4133 1-A (holotype: MICH) 

Paratype: new GUINEA. Morobe Province: Boana, 1,219 m, 4 Jul 1940, M.S. Clemen 
41330 iUlCH). 

ned for the indefatigable collector Mary 

Fig. 3. Fittingia manae Stone (drawn from holotype). A-D. Floi 
with pedicel and part of partial inflorescence with bracts. B. Ir 
showing densely papillate inner faces of corolla-lobes, and short 
ginate anthers. C. Stamen, abaxial view; note the non-glandular 
discoid stigma. E. Diagram of inflorescence. F. Leaf 

Stone, Malesian Myr 

Fig. 5. Tapeinospcrmcjinpes^um^. Phc 

Stone, Malesian Myrsinaceae 271 

Strong Clemens who with her husband Joseph, and later alone, obtained 
numerous plant collections in the Philippines, Borneo, and New Guinea. 

Notes: The dioecious habit, 4-merous flowers, and axillary inflorescences 
indicate that this species belongs to the genus Fittingia. Among other gen- 
era of the family that occur in New Guinea only Loheria could be consid- 
ered as a possible alternative, but that has short lateral shoots with flowering 
panicles subtended by a subinvolucral group of bracts. In the absence of 
pistillate flowers, some doubt may remain, but the general habit and slen- 
der panicles, sharply acuminate leaves and low habit suggest Fittingia much 
more than Loheria. In Fittingia also the corolla seems to lack internal 
papillosity, while it is common in Loheria. 

Among the species oi Fittingia, F. mariae may be readily distinguished 
by the slender, sharply acuminate leaves and very slender panicles. Collec- 
tion of pistillate materials including fruits remains a desideratum. 

Tapeinosperma Hook.f. 

Sleumer (1988) lists four species oi Tapeinosperma in New Guinea. Among 
the L.J. Brass collections the following new species has been identified. 

Tapeinosperma filipes B.C. Stone, sp. nov. (Fig. 5). 

Arbusculaplerumquemonocaulosa usque ad 1-1.5 malta; caulisap 

icem versi 

elongatis teret.bus 30-35 cm longis. Innovationes minute ferrugmeo-t< 
0.06 mm longis capitatis cellulo apicale anguste ovoideo vel lanceoloic 


mfra glabrata (costa media excepto); costa supra basi anguste canalicul 

lato, infra 

) minute tomentellus demum glabratus. Ramuli fertili tereti laevi 1-2.5 mm diar 
S cm longi (usque ad foliam primam); foliis reductis 7-1 1 cm longis subulatis; a: 
;is pedicellisque minute fermgineo-tomentellis. Inflorescentia terminalis. Flores n- 
i subglobosi, pedicellis 2.5-3 mm longis; calyce cupulato 5-lobato, basi sparsiter hi 

)-punctatis, integris. Stamina ca. 1 .3 mm longa, filamento 0.2 mm longo, antheris obi 
, subacutis, dorso sublaeve vix glanduloso vel glandulis obscuris singulis in conm 
s. Ovarium ca. 0.5 mm longum, glabrum, sensim in stylo 0.7 mm longo produ( 
ate minute truncato obscuriter discoideo et minime papilloso. Placenta ca. 0.4 mn 

Known only from this collection from lower m 
Etymology: The epithet, filipes, refers to the \ 

like inflorescence axes (fila = thread; pes = foot, stalk). 

Notes: This distinctive species was cited as an indeterminate Ardisia by 
Sleumer (1988: 103), but the discoid-truncate stigma and strongly reduced 
ovular number exclude it from that genus. The flowers are very small but 
seem to be hermaphroditic, and in this respect conform to the condition in 
Tapeinosperma. The very long slender leaves and extremely slender open 
panicles borne on long, slender axillary branchlets bearing 1 or 2 reduced 
leaves are further characters of this species. 

Sleumer, H. 1988. The genera Dtscocalyx Mez, Fittingia Mez, Loheria Mei 

Hook.f. (Myrsinaceae) in New Guinea. Blumea 33:81-107. 
Stone, B.C. 1989- Myrsinaceae. In: Ng, F.S.P. (ed). Tree flora of Malaya. ^ 
Stone, B.C. 1990. Studies in Malesian Myrsinaceae V. Additional new s| 
Proc. Acad. Nat. Sci. Philadelphia 142:21-58. 






Department of Botany 

University of Texas 

Austin, TX 7 87 13, U.S. A. 

1 Symphorkarpos have revealed the existe 
)m the Cuchumatanes Highlands of Gu; 
le species oi Symphorkarpos from Mexico 

Recientes estudios de los Symphorkarpos mexicar 
de una nueva especie, que se describe e ilustra a c 
de Guatemala. Ademas, se ofrecen claves y mapas 
de Mexico y Guatemala. 

Key Words: Symphorkarpos, Caprifoliaceae, Guatemala, Mexico. 
SymphoricarposguatemalensisJ.K. Williams, sp. nov. (Fig. 1) 

Symphorkarpj mtcrophyllo HBK. simiUs sed differt fauce corollarum glabra et floribus ac foliis 

Erect shrub, 2 m tall, much-branched, young twigs strigose. Leaves oval 
with rounded apices, tapering toward base, 3-6 mm long, 2-3 mm wide, dark 
green above, margins entire, glabrous to slightly villous, coriaceous; petioles up 
to 1 mm long. Flowers solitary or in pairs; pedicels up to 0.5 mm long, with 
two apical bracteoles 2-3 mm wide; calyx glabrous, 1-2 mm long, irregularly 
5-toothed; corollas white, narrowly campanulate, actinomorphic, papillate to- 
wards base within, 4-6 mm long, tubes 3-5 mm, lobes 1-2 mm, erect; stamens 
5 , epipetalous, barely exserted, anthers 1 .0-1 . 5 mm long, ca. half the length of 
filaments; style glabrous, 3^ mm long, stigma capitate. Fruit a globose berry, 
6—7 (10) mm long, white, and tipped with persistent calyx. 

Distribution and Ecology: Limestone ridges of the Sierra de los Cuchuma- 
tanes in Guatemala (Fig. 2), 3000-3500 m; flowering July to September. 
^UEHUETENANCK): 3 mi SW of San Mateo, Ixtatan, 5 Aug 1965, D.£. 


I. Symphamarpoi gua tenia letisi 

lined: GUATEMALA. Huehuett 


/ (F); 1 

ma 30022 

\ MO); betwee 

1 Llanos San Miguel, road t< 
xoy, 17 Nov 1967, Molina 21248 (F); dry slopes of El Mirador, 12-23 Jai 
2a 16303 (F); along road to San Pedro Soloma, 3 mi SW of San Mateo Ixtatan 
, BreeJlope 11544 (F); limestone ridge near Chemal, Sierra de los Cuchumatanes 
32, L.O. Williams 21979 (F); just below Calveras, 29 Nov 1962, LO. William 
5ierra de los Cuchumatanes, at KM 31 1 on Ruta Nacional 9N (between Paqui: 
1), small limestone ridge ^'it'a j uniperm standleyi, 2 Aug 1959, Beamati 298^_ 
r Tunima, Sierra de los Cuchumatanes, 6-^7 Jul 1942, Steyermark 4SJ()4 (F) 
in region of Chemal, 31 Dec 1940, Standley 81624 (F); rocky ridges beneatl 
1 Cuchumatanes, 15 Sep 1934, Skutch 1253 (F). 

Williams, Symphoricarpos guatemaL 

area) and S. mkrophyllm (square 

This narrow endemic of the Sie 
subgenus Anisanthus G.N. Jones ( 
lobes shorter than the corolla tul 
guatemalensis is most similar to S. t, 

L de los Cuchumatanes is a member of 
les 1940), as evidenced by the corolla 
and its included style. Symphoricarpos 
^ophyllus but differs in its glabrous (vs. 

villous) corolla throat, dark brownish-green leaves (vs. light green), and 
leaves and flowers of half the dimension. The leaves of ^. guatemalensis are 
3-6 mm long and 2-3 mm wide, compared to those of 5. microphyllus which 
are 10-25 mm long and 7-15 mm wide (Fig. 3). In addition, the corolla 
length of ^. guatemalensis is 4-6 mm, compared to 9-1 mm in S. microphyllus 
(Fig. 1). 

In his monograph, Jones (1940) reported that species oi Symphoricarpos , 
other than S. microphyllus, are rarely found in Mexico and further south. 
Accumulating collections from Mexico, however, have shown that the ge- 
nus IS more prevalent there than once suspected. In order to place the new 
species in a broader perspective and to provide a preliminary taxonomic 
evaluation of the Mexican and Central America species, I have constructed 
a key to the species oi Symphoricarpos in this area based on specimens housed 

at BRIT, F, LL, and TEX. Characters used to delineate taxa have been es- 
sentially adopted from Jones (1940) and Gray (1873). 

Symphorkarpos is divided into two subgenera: subg. Symphorkarpos , char- 
acteri2ed by an exserted style and corolla lobes longer than the tube, and 
subg. Anisanthus, characterized by an included style and corolla lobes shorter 
than the tube. There is only one member of subg. Symphorkarpos, S. 
orbkulatus, native to Mexico. All other species in Mexico and Central America 
belong to subg. Anisanthus. 





an tube; fru. 

its red 

I S orbtadam 

, style aiway 


m long, glabrous w.rh 


2 S longtflorus 

5 mm wide; 


Lrchmg, 1-] 


- branches 

.entire... 4. S.mkrophy/l//s 
leaves of older 


Symphoricarpos orbiculatus Moench 

This is the only member of the subg. Symphoricarpos native to Mexico; other 
species of the subgenus occur throughout North America and Asia. This par- 
ticular species is unique to the genus in having red fruits (Fig. 5). 

Representative specimens: MEXICO. Coahuila: Muzquiz, along Sabine River, 1 1 Jul 1936, 
Marsh 475 (TEX). Nuevo Leon: Monterey, Diente Canyon, 20 Jul 19.30, Mueller 103 (F, TEX). 

Symphoricarpos longiflorus A. Gray 

This species is distinguished by its salverform corollas 1 1-13 mm long, the 
largest of any corolla among the Symphorkarpos of Mexico. Other features that 
characterize this species are its sessile anthers and solitary basal nectary (all other 
Mexican taxa have five nectaries). Another feature which helps separate this 
species from other Mexican Symphorkarpos is a pubescent style; this is a plastic 
character, however, and not always reliable. The foliage and style of populations 
in Mexico range from completely glabrous (in Baja California) to pubescent (in 
northern Chihuahua) (Fig. 4). 


itative specimens. MEXICO. Baja California Norte: Sierra San Pedro Martir, 


slope c 

)f Cerro de la Culpa, 31° 02'N, 115° 27'W, 27 Jul 1970, Moran 17890 (TEX). 



l'2866 (TEX). 

Symphoricarpos microphyllus HBK. 

Symphorkarpos microphyllus is the largest of the Symphoricarpos species, some- 
times attaining a height of over 2 meters. It is readily distinguished from S. 
palmeri and S. parishii in its erect habit and by having all its leaves entire (Fig. 3). 
This is the most widespread Symphoricarpos species in both Mexico (Fig. 4) 
and Guatemala (Fig. 2). Throughout most of its range S. microphyllus is pubes- 
cent, but there is a rather large isolated population of glabrous individuals 
in the Sierra Madre Oriental along the border of southern Coahuila and the 
Midwestern Nuevo Leon border (Fig. 4). 

Representative specimens. GUATEMALA. Chimaltenango: slopes of Volcan de Acatenango, 
above Las Calderas, 3 Jan \9i9,Standley 61833 (F); Volcan deAgua, 22 Jul 1951 J.R.Johnston 
9iO (F). El Progresso: between Calera and summit of Volcan Siglo, 21 Jan l9A2,Steyermark 
43042 (F). Huehuetenango: across river from San Juan Atitan, 8 Sep 1942, Steyermark 52018 
(F); Cerro Pixpix, above San Ildefonso Ixtahuacan, 15 Aug 1942, Steyermark 50566 (F). 
Quezaltenango: Summit of the Sierra Madre Mts, 15 km N of Ostuncalco, 9 Dec 1963, L.O. 
Williams 25536 (F); Sierra Madre Mts, 5 km N of Ostuncalco, 8 Dec 1963, L.O. Williams 
25472 (F); Cuesta de El Caracol, Sierra Madre Mountains, 5-8 km N of San Juan Ostuncalco, 
department of Quezaltenango, 1 1 Dec 1962, WilHams 22768 (F, TEX); mts SE of Palestina, 
along new road to San Juan Ostuncalco, 2 1 Jan I94l, Standley 84354 (F); Volcan Santo Tomas, 
22 Jan 1940, Steyermark 34804 (F); Volcan Santo Tomas, 22 Jan 1940, Steyermark 34804 (F); 
Volcan Santa Maria, between Santa Maria de Jesus, Los Mojadas, and summit of volcano, 1 2 Jan 
1 940, Steyermark 34050 (F); slopes of Volcan Santa Maria, above Palojunoj, 6 Mar 1939, Stand- 
ley 67668 (F); mts above Olintepeque, 20 Feb 1939, Standley 65956 (F); Volcan Santa Maria, 
27 Jul 1934, Sk/ach 870 (F). Sacatepequez: Slopes of Volcan de Agua, above Santa Maria de 
Jesus, 1 1 Feb 1939, Standley 65108 (F). San Marcos: between Sibinal and summit of Volcan 
Tacana, 19 Feb 1940, Steyermark 36080 (F). Totonicapan: ravines in Sierra Madre mts near 
VillaLasCruces,8kmSWofTotonicapan, 13Dec 1963, L.O. VZ/Y/z^m 22969 (F); Nebuloso de 
Maria Tecun, 21 Nov 1965, Molina 15886(F). 

MEXICO. Chiapas: Zinacantan, 6 Oct 1986, Ton 9333 (TEX); San Juan Chamula, 2 Jun 
1 988, Ruiz 883 (TEX); Valley floor of Zinacantan Center, 1 3 Apr 1 966, Laughlin 668 (F); slope 
with Oak and Pine, along road to Zontehuitz near Piedrachitas, 25 Aug 1964, Breedlove 7127 
(F); near Ik'al Nab lake, on the boundary between Chamula and Zinacantan, along road to 

along road to Oxchuc and Ocosingo, 21 Sep 1972, Breedlove 277 50 (F). Coahuila: Sierra El 
Coahuilon, Arteaga, 17 Jun 1991, H/Kf<?« 20992 (TEX); Sierra Pilote de Fierro, 25° lO'N, 
101° 25'W, 24 Oct 1991, Carranza 1221 (TEX); Las Vigas canyon. Sierra de Arteaga, 25° 
20'N, 100° 39' W, 5 Jun 1987, Villarreal 3739 (TEX); Sierra la Marta E of Cerro Moro, 22 Jul 
19^5, Gtnzburg 141 (TEX); Muzquiz, Sabmas river, 1 1 Jul 1936, AI^rj/>47^ (BRIT-SMU). 
Distrito Federal: volcanic soil, SE side of D.E, 12 Jul 1947, Barkley 2215 (TEX); Desierto de 
los Leones, 5 Nov 1942, Steyermark 52250 (F). Durango: canyon head, 5 1 mi SW of Durango, 
13 Aug 1957, Waterfall 13727 (BRIT-SMU, F, TEX). Guanajuato: San Miguel de las Casitas, 
8 Sep 1990, Ventura 8692 (F). Hiidago: on Hwy 104 NE of Pachuca, 49-7 mi E of Ajacuba, 
10 Jul 1985, a/i^^« ^493 (TEX); San Vicente, 16 Aug 1937, F;j/^er 3 7i9i (F); upper cliffs 

near Metepec Station, 27 Jun 1904, Prtngle 13010 (F); Sierra de Pachuca, 20 Jul 1901, 
Pnngle 9480 (F). Jalisco: Nevado de Colima, brecha entre El Izote y la Micronda las Viboras, 
5 Feb 1992, Chazaro 6868 (TEX); NE slopes of Nevado de Colima, below Canoade Leoncico, 
10 Sep 1952, McVaugh 1287^ (BRIT-SMU). Mexico: 1 1 mi E of Amecameca, 31 Jul 
1975, LeDoux2037 (TEX); between Toluca and Mexico, 24 Jul 1962, Moheeci 39 (TEX); 
pine forest of Las Crucea, 12 Jul 1932, Hinton 1013 (F); 2 km SE of Totoloapan, 18 km E 
of Tezcoco, 10 Jul 1979, Koch 7949 (F); valley in mountains, 16 mi E of Toluca, 18 Aug 
1957, Waterfall 14067 (F). Michoacan: on road to Uruapan, 4 mi S of Cheran, 3 Nov 
l958Jones 23326 (TEX), hills of Patzcuaro, 1 1 Oct 1892, Pr;V^ 426/ (F); Cerro Huashan, 
2 km S of Nahuatzen, 28 Jun 1990, Garaa 2724 (F); Zitacuaro, 19 Aug 1938, Hwton 
13120 (F); 1 km W of Rancho Seco on Hwy 37, 2 km W of Morelos, 13 Sep 1986, Swage/ 
167 (F); 6 km SE of Villa Madero, 30 Sep 1982, Nunez 4732 (F). Morelos: Tepoztlon, 15 
Jul 1938, Kenoyer A340 (F). Nuevo Leon: 6 km SE of La Trinidad, summit of one of the 
picachos of Sierra Cebolla, 25° ll'N, 100° 07'W, 5 Aug 1988, Patterson 6174 (TEX); 
SierraLaMarta,Galeana, 6Sep 1981, Hinton 18343 (TEX); Cerro, El Potosi, 21 Jun 1969, 
Htnton 17115 (TEX); 1 .5 mi SW of Pabillo, 21 Jul 1 958, Correll 19933 (TEX); Municipio 
Galeana, open pine forest near Cerro Potosi, 23 Jul 1938, Schneider 933 (F); peak of Cerro 
Potosi, Municipio de Galeana, 21 Jul 1935, Mueller 2236 (F); Sierra Potosi, by the N 
hogback, 20 mi NE of Galeana, 26 Jul 1934, Mueller 1243 (TEX); Lerios Mts., Coahuila, 
45 mi E of Saltillo, 10-13 Jul 1880, Palmer 390 (F). Oaxaca: Cerro Verde, Jul 1908, 
Purpus3318{¥). Puebla: El Cimarron, 28 Jul 1971, Vetera 3954 (TEX); el Chamizal, 20 



, Barkley 2469 (F). Queretaro: 0.5 km of Cuatro Palo< 
989, Carranza 21 10 (F); Campo Alegre, 3 km S of Sa 
3298 (TEX). San Luis Potosi: 7 km SW of Pozueln 

Symphoricarpos palmeri G. N. Jo 

Symphorkarpos palmeri is a relativel 
Trans-Pecos region of Te? 

Mexico. It is recognized by its trailing habit and crenate-mucronate leaves 
of both flowering and non-flowering branches (Fig. 3). The species has mostly 
pubescent stems and leaves, but individuals with glabrous stems occur 
throughout its range (Fig. 5). 

^ of Cerr 

, El Ja, 

in, 29° 06'N, 



cacho del Centinela, 24 Aug 


Symphoricarpos parishii Rydb. 

Symphoricarpos parishii is distinguished from the other Mexican taxa by 
its trailing habit, entire leaves of the flowering branches, and lobed leaves 
of the older non-flowering branches (Fig. 3). Specimens studied from Mexico 
have glabrous stems, although pubescent individuals are found scattered 
throughout its range in the United States. This species has been regarded 
as a variety of both S. oreophilm A. Gray by Cronquist et ai. (1984) and S. 
rotundif alius A. Gray by Dempster (1992), but this problem is beyond the 
scope of the present paper (Fig. 5). 

Fig. 5. Distribution o 

<'. parish! i (square) in Mexi 

Representative specimens. MEXICO. Baja California Norte: Parque Nacional Sierra 
San Pedro Martir, Vallecitos, 31° 02'N, 115° 27.5'W, 1 Sep 1985, Thome 61457 (TEX); 
Sierra de Juarez, Parque Nacional Constitucion de 1857, Laguna Hanson & adjacent rocky 
flats and slopes, 26 May 1987, Thome 62362 (F); upper east end of Vallecitos meadow, 31° 
OO'N, 115°27'W, 3Jul 1973, Mom« 27082 (TEX). 


I would like to give special thanks to the very warm and generous Dr. Guy 
Nesom for suggesting this project to me, as well as for the Latin diagnosis and 
many fine readings of the manuscript. Nancy Webber was beneficial in provid- 
ing beautiful and expedient illustrations. Alan Prather and Dr. B. L. Turner 
provided helpfol comments. Cooperation from BRIT-SIMU, F, and TEX-LL was 
also much appreciated. 

Dempster, L.T. 1992. A nomenclatural change mSymphoricarpos. Madr 
Gray, A. 1873. Revision of the genus Symphorkarpos.]. Linn. Soc, Bot 
Jones, G.N. 1 940. A monograph of the genus Symphorkarpos. J. Arnol( 



The Tucker Center 

College of Forestry 

Stephen F. Austin State University 

P.O. Box 6109 

Nacogdoches, TX 75962-6109, USA. 

•s pool refers t( 
istern North t 

It of regional species 
n ecologically deter- 

cle extincion y especiacion bajas, mientras que la gran riqueza en el noroeste de Norte America 
es debida a especiacion reciente rapida. Los datos no explican la dominancia teorica del 
determinismo ecologico en favor de la historia geofisica regional especifica como base para 
explicar la diversidad regional de especies. Se propone un "pool" de especies como resultado de 

Species diversity is a central theme of environmental studies. Scientists have 
accumulated much data on species diversity over the past 250 years. However, 
there are still no answers to some basic questions. One critical question that is 
frequently asked by scientists is why there are just so many species in a region, 
and not more or less than that number. In other words, what factors determine 
regional species richness? In fact, a comprehensive, rigorous, and general theory 
of species diversity is lacking (Solbrig 1991). Ricklefs (1987) thus claimed that 
species diversity is "the major, unexplained pattern in natural history," one that 
"mocks our ignorance." 

One dominant hypothesis is that available energy determines regional spe- 
cies richness (Hairston et al. I960, MacArthur and Wilson 1967, Carrie and 
Paquin 1987, Moore 1987, Adams and Woodward 1989, Andersson 1990, 
Currie 1991). Currie and Paquin (1987) and Currie (1991) clearly claimed that 

regional tree species richness can be explained in terms of actual evapotranspira- 
tion (AET). Alternatively, White (1983) claims that the tree species diversity 
pattern of the Northern Hemisphere is the product of extinction. Goldsmith 
(1985) believed that periodic, or episodic, catastrophic destruction of life was 
caused by comet or asteroid impacts. However, there are three critical problems 
with these positions: First, almost all studies of species richness concentrate, 
very naturally, on just one taxonomic group (e.g. pines, oaks) (iVfoore 1987). But 
each group may have some peculiar requirement of its own that obscures the 
general causes of species diversity (Moore 1987). Second, studies were often 
made using small biogeographic regions. Micro-environmental variations greatly 

tantly, the conceptual definition of a species has not been consistent from tax- 
onomist to taxonomist nor from region to region. 

To minimize the observation biases caused by the factors mentioned above, 
this study examined all seed plants in the larger biogeographic regions in the 
Northern Hemisphere. Eastern Asia in this study refers to the eastern forest 
region of China, Far East and eastern Siberia of Russia, and Korea (Fig. 1). 
North America includes the continental regions of Canada and United States. 
The forests of North America are naturally divided into those west and those 
east of the Great Plains (Fig. 2). The boundary of the boreal zone in the North- 
ern Hemisphere follows Larsen (1980) with modification in eastern Asia. The 
boreal region in Europe includes the area westward from the Ural Mountains 
(Fig. 3: 1). Boreal eastern Asia includes eastern Siberia in Russia and the Da 
Xingan Ling Mountains of China (Fig. 3: II). The east-west division of the 
boreal zone in North America follows Daubenmire (1978). The eastern section 
includes the area east of the Rocky Mountains (100° W) (Fig. 3: IV) and the 
western or Cordilleran Section extends westward (Fig. 3: III). 

In eastern Asia, the temperate forest region includes the Xiao Xingan Ling, 
Wanda, and Changbai mountains in Northeast China, southern portion of Far 
East of Russia, and northern Korea (Fig. 3: V). This region is known as the 
Korean pine conifer and hardwood forest region (Chou and Li 1 990). In western 
North America, the temperate forest region covers portions or all of the Pacific 
Northwest states of Oregon, Montana, Idaho, Washington, and southern Brit- 
ish Columbia (Fig. 3: VI). This region includes the Pacific Northwest forests 
(Franklin 1988) and the northern portion of the forests of the Rocky Mountains 
(Peet 1988). In eastern North America, the temperate forest region extends 
from northern Minnesota upper Great Lakes region and eastward to southern 
Canada and New England, including, toward the southeast, much of the Appa- 
lachian Plateau in New York and northern Pennsylvania (Braun 1967, Fig. 3: 
VII). The region is known as the Hemlock-white pine-Northern Hardwoods 
region (Braun 1967, Vankat 1 979) as well as the mixed conifer and deciduous 
forest region (Bailey 1976). 

The main data sources of this study include field investigations, specimen 
)bservations, personal communications, and literature (including flora and fos- 
il records). The primary data for eastern Asia were collected fi-om a comprehen- 
ive forest survey conducted by authors and their colleagues at Northeast For- 
estry University and numerous forestry agencies during 1984-1988. Data for 

North America is based on literature, supported where possible, b\ field collec 
tions, herbarium investigations, and personal interviews during 1 9H9- 1 992 B> 
using established methods and the same principal ubscivci jn the hciburium 
and field, the observational biases that often plague broad-scale comparisons 
were minimized. 

Li and Adair, Specie 


v^ ^'\ T 

7 7' /' / 










160 /^ 

^^j^::';5 ?y^X^^^ 









\ X 







1,20- no- ,00- 9 


Fig. 3. Map showing the boreal and temperate forest 
America; V-VII — temperate forest zone: V-eastern A 

Five sheets of voucher specimens of each t 
boreal and temperate forests of eastern Asia we 
tigations. The location, life history, growth forms, fiinction of woody plants, 
vegetative organs, habitats, and other characteristics such as flower color of each 
species were recorded for each specimen. The voucher specimens were retained in 
the Institute of Plant Sciences at Northeast Forestry University in Harbin, China. 
Manuals involving the floras of East Siberia, Far East, Korea, and Japan were also 

used to identify voucher specimens. Herbarium collections, located in Northeast 
Forestry University, the Botanical Institute in Beijing, and the Applied Ecologi- 
cal Institute in Shenyang were also used to confirm identification of voucher 
specimens. Seed plant checklists of the boreal and temperate forests of eastern 
Asia were created separately from these comparisons. Additional floristic check- 
lists were created separately for boreal Europe, boreal western North America, 
boreal eastern North America, temperate Japan, temperate western North America, 
and temperate eastern North America using scientific references: Ohwi (1965), 
Polunin (1959), Gleason and Cronquist (1991), Hitchcock and Cronquist (1974), 
and Scoggan (1978-1979). 

In both boreal and temperate forest comparisons of Europe, eastern Asia, and 
western and eastern North America, floristic analyses were made at the family, 
genus, and species levels. Evolutionary analyses of flora were based on the diversi- 
fication ratios (family and genus size), diversification rate (Eriksson and Bremer 
1992), and flower characters. Phylogenetic analyses were made in terms of three 
categories: 1) primitive — subclasses Magnoliidae and Alismatidae; 2) intermedi- 
ate — subclasses Caryophyllidae, Hamamelidae, Dilleniidae, Rosidae, Arecidae, 
Commelinidae, and Zingiberidae; and 3) advanced — subclasses Astendae and 
Liliidae (following Cronquist' system in 1988). 


The great plant species diversity of eastern Asia is weU-known and has been 
widely emphasized by scientists (Sargent 191.3, Hu 1935, Takhtajan 1969, Li 
1952, Wu and Wang 1983, Boufiford and Spongberg 1983, Latham and Ricklefs 
1992). To date, 25,480 species of native vascular plants have been identified in 
eastern Asia, representing 300 families and 2,875 genera (Li 1993). This amounts 
to 10.7% of the total known species in the world and about 1.6 times the figure 
for North America (Table 1). The great plant diversity in eiistern Asia is the 
product of long evolutionary history. The geological and palcobotanical records 
failed to support the existence of continental Quaternary glaciers in most of 
eastern Asia and thus extinction is not a main process in the development of the 
extant eastern Asian flora. Also, eastern Asian flora consists of a host of taxa that 
are presumed to be phylogenetically primitive, with many occurring as mono- 
typic taxa in the subtropical or tropical regions. These factors have led to the 
concept that eastern Asia was the evolutionary source of modern flora in the 
Northern Hemisphere (Takhtajan 1969, Latham and Ricklefs 1992). 

Ferns are phylogenetically primitive vascular plants. 52 families (80.0% of 
the total flora of the world), 204 genera (46.0%), and 2,300 species (19.5%) of 
ferns are known in eastern Asia. The family, genus, and species numbers are 3.5, 
3.4, and 6.7 times those in North America, respectively Gymnosperms, another 
phylogenetically primitive group of plants, are represented by ten families 

d on Checklist of D.E.] 

in eastern Asia, but only five in North America. Eastern Asia contains almost all 
primitive monotypic families, including Ginkgoaceae, Cycadaceae, and Gneta- 
ceae. Ginkgo, for example, is the oldest of all trees, it dates back to the Triassic 
some 200 million years ago (Flora of North America Editorial Committee 1 993). 
It is now naturally restricted to China although it was quite widespread in the 
world during the Cretaceous (Hsu 1983). Conifers apparently represent the 
greatest taxonomic diversity in eastern Asia. They are distributed among Eu- 
rope, eastern Asia, and western and eastern North America, respectively, ap- 
proximately in the ratio 1.0 : 3.4 : 2.1 : 1.6 by genus and 1.0 : 5.9 : 2.4 : 1.2 by 
species (Table 2). 

There are 2,637 genera and 23,000 species of angiosperms belonging to 238 
families native to eastern Asia (Li 1993). The family, genus, and species num- 
bers in eastern Asia are 1 .4, 1 .2, and 1 .6 times those in North America, respec- 
tively. Almost all ancestral families of angiosperms based on fossil records have 
greater diversities in eastern Asia than in North America. Chloranthaceae, as 
the earliest angiosperm (appeared in the early Cretaceous, MuUer 1981) has three 
genera and l6 species in eastern Asia and only one genus and one species 
in North America. Aquifoliaceae, one of the earliest woody angiosperm families 
(Muller 1981) represents 168 species in eastern Asia and about 20 species in 
North America. Moreover, iVlagnoliaceae, the most phylogenetically primitive 
woody family of angiosperms (Hutchinson 1973), contains 12 genera and 250 
species ranging from Asia to North America (Wu and Wang 1983). Ten genera 
and 100 species of this family are known in eastern Asia, but only two genera 

nd 9 species are known in North America. Hamamelidaceae, another primi 
ive family, has 25 genera and 90 species widely distributed in eastern anc 
DLitheastern Asia and North America with some reaching south to Austra 
a and Africa. 17 or 18 genera and 70 species of this family are known ir 
astern Asia. Disanthus, Exbucklandria, and Rhodoleia, ancestral genera o 

Li and Adair, Species pools 289 

the family (Takhtajan 1969), are all found in southern China. Only three 
genera and five species are present in North America. 

The data above imply that eastern Asian flora has a long evolutionary 
history. However, the analysis of diversification rates (Eriksson and Bremer 
1992) of families based on fossil records shows that species diversification 
rates* of most families in eastern Asia are much lower than those in North 
America. 20.7% of the total species in eastern Asia belong to families with 
low diversification rates (R<0.15 my'^), while only 11.2% of the total spe- 
cies in North America belong to the same families. 

Thus, we concluded that the great taxonomic richness of eastern Asia is largely 
the result of a lengthy evolutive history coupled with low extinction and low 

Geologically, northwestern North America is young. But the northwestern 
North American flora has the highest species diversity among all northerly re- 
gions in the Northern Hemisphere (Table 3 and 4). This great species diversity 
results from high speciation in a short evolutionary history. 

The taxonomic and evolutionary structure of extant flora is consistent with 
the concept of a geologically young western North America. In the temper- 
ate zone, for example, western North America has a total 3,161 species, which 
is about 50% of the total species of all three temperate floras and about 1.5 
times that of either eastern Asia or eastern North America. However, this 
flora has 19 families fewer than in eastern Asia and 27 families fewer than in 
eastern North America (Li 1993). Therefore, families of temperate floras, on 
average, have more species in western North America (roughly 30 species per 
family) than in eastern Asia (roughly 17) and eastern North America (15, see 
Table 5). Theoretically, the larger the family (containing more species), the 
more evolved (Stebbins 1981), because family is parental to species. From 
this point of view, the temperate flora in western North America is young. 

The evolutionary analysis of flowers strengthens this statement. The solitary 
flower, both terminal and axillary, is the initial form of arrangement for flowers, 
while inflorescence is more advanced and of a greater biological advantage 
(Takhtajan 1991). In the temperate zone, western North America contains the 
highest proportion of inflorescence (87.8% of the entire flora, compared with 
80.4% in eastern Asia and 79T% in eastern North America) (Li 1993). Flower 
color is an important factor affecting pollination of plants. Green flowers are 

largely pollinated by wind, and thus the anemophilous plants are claimed to be 
primitive in evolution (Takhtajan 1991). Alternatively, bright colored flowers 
are usually pollinated by insects, and entomophilous plants are considered more 
advanced m evolution. In western North America, only 21.8% of the total 
species have green flowers, a lower percentage than in either eastern Asia (27.8%) 
or eastern North America (33-6%). However, colorful flowers (white, yellow, 
pink, rose, purple, blue, and red) are more frequent in western North America 

3 Adair, Species pools 

and comprise 78.2% of the total flora. The same colors account for 72.2% in 
eastern Asia and 66.4% in eastern North America. 

Life history, as the result of natural selection, reflects the long-term adapta- 
tion of a plant to its environment. Annuals are unknown in primitive ferns and 
gymnosperms, but their sexual reproduction processes enable them to persist in 
habitats that restrict the establishment of perennials. Therefore, annuals are 
believed to arise relatively late in the evolution of terrestrial plants (Bazzas and 
Morse 1991). Annuals are distributed among eastern Asia, western and eastern 
North America at the ratio of 1.0 : 2.5 : 1.0. Alternatively, as the early evolved 

America at ratios based on total numbers of both genus and species as follows: 
1.3 : 1.0 : 1.9. Obviously, the temperate flora in western North America is 
relatively young in evolution relative to the other two temperate floras. 

Further, phylogenetic data strengthen the argument. Magnoliidae and 
Alismatidae are the most primitive taxa of flowering plants in Cronquist's system 
( 1 988). They, together with gymnosperms represent the lowest diversity in tem- 
perate western North America (16 families and 56 genera) and the highest in 
rn Asia (23 families and 72 genera). But Asteridae and Lihidae, 
t advanced taxa in Cronquist's system have the highest diversities in tem- 

perate western North America (243 genera, compared with 233 genera in tem- 
perate eastern Asia and 199 in temperate eastern North America respectively). 

In brief, all these evidences from taxonomic structure of flora, flower diver- 
sity, life history and phylogeny show that northwestern North American flora 
is relatively young in evolution. However, data on diversification rates (R) for 
families show that there is high species diversification in northwestern North 
America. A total of 25 families with low diversification rates (R<0.10 my"') 
are recorded in the temperate regions, and they are distributed among eastern 
Asia, western and eastern North America, respectively, at a species ratio of 2 : 1 
: 3 (Table 6). Western North America clearly has the smallest taxon diversity 
among families with low diversification rates. In contrast, the nine families 
with higher diversification rates (R>0.60 my"^) are distributed at a ratio of 
total number of species of 1 : 3 : 1 in eastern Asia, western and eastern North 
America. Further, three genera with the highest diversification rates, Astraga- 
lus, Senecio, and Carex display maximum species richness in western North 
America and exceed that of any other region (279 species relative to 51-145 
species in the temperate regions and 144 species relative to 56-90 species in the 
boreal regions). Boreal floras show a similar pattern (Li 1993). 

Thus, there is little doubt that the data indicate that the northwestern 
North American flora is the product of high species diversification and rela- 
tively recent evolution. Considering that the species-rich northerly flora of 
western North America has over 50% of the total species restricted to this 
region, we conclude that this great species richness in northwestern North 
America is largely the effect of rapid speciation rather than immigration. 

a given time. This regional species pool is a relatively static spot in a dynamic 
river of species diversity development over time. It is the result of two additive 
processes: speciation and immigration; and, one subtractive process: extinction 
(both locally and globally) (Fig. 4, Table 7)! Thus, a species pool has largely an 
historical rather than an ecological basis for its existence (Li 1993). 

Speciation is the formation of new species from pre-existing ones usually 
by a process of improved adaptation of survivors to the environment. New 
species may be better adapted to the environment and replace the less well 
adapted ones. Speciation began long ago and is still going on. It is best 
characterized as a regional subset of the global genetic pool. The rate of 
speciation depends not only on environmental diversity within the pool, 
especially under conditions of geographic or ecological isolation, but also 
on chance. Thus, it is impossible to predict the speciation rate for a region. 

Immigration, however, does not involve global genetic variability, but does 

Li and Adair, Species pools 


Species Number 


Adaptation to environments 

Global Local 
Failed adaptation to environments 

Fig. 4. Speci 

me (After Li 1993). 

contribute to regional changes. During migration, species may evolve. If generic 
change makes it possible to distinguish new species from the parent species, this 
process is evolutionary (speciation); if not, it is called immigration. Both natural 
selection and chance play roles in immigration of species to a region. This also 
means that the absence of a species in a region may result purely from chance 
rather than deterministic natural processes. 

Extinction of a species can occur locally (exclusion or emigration), in which 
one or more populations vanish but others survive elsewhere, or globally, in 
which all members of the species population everywhere in the world vanish. 
Extinction of a species is a natural process. The fossil record indicates that 
most, if not all, species have a finite life span, averaging between 1 and 10 million 

Li and Adair, Species pools 2 9 5 

years (Solbrig 1991). Drastic environmental change is a major cause of species 
extinction especially on a global scale. Because the environment is in a constant 
state of transformation, some species are always being lost while others are added. 
Some changes in the physical environment are cyclic and periodical, while others 
are less predictable. Thus, it is impossible to establish a normal rate of species 
change and based on the data presented in this study, it may be impossible to do 

A species pool has the following main characteristics. 
Liquidity and Irreversibility 

A species pool is a dynamic flow of species through a storage area. Adapted to 
new environments, species that have become locally extinct (exclusion) may re- 
migrate back into a region. However, the natural process from speciation to 
extinction is irreversible. Species that have become globally extinct cannot be 
recovered naturally. Thus, conservation of biodiversity is needed on a global 
level. But, it is impossible for humans to interrupt the natural extinction pro- 
cess and store all species on a global scale. Man can temporarily store as 
many species as possible in regional species pools, but even these are present 

A species pool is not the accumulation of species only over long evolutionary 
time spans, and species-rich communities may not always be the oldest as 
Whittaker (1977) has stated. Western North America has a young geological 
history. The rapid rise of the Rocky Mountains in the Tertiary and extensive 
glaciation in the Pleistocene caused widespread extinction of life in western 
North America. However, western North America has the greatest species pool 
in the northerly regions of the Northern Hemisphere. The northerly flora of 
western North America, which is both young and species-rich, is a striking 
example that fails to support Whittaker's hypothesis. This fact is also not con- 
sistent with White's hypothesis that emphasizes that the differences in taxon 
richness among eastern Asia, North America, and Europe results from differing 
Pleistocene extinction rates in the three areas (White 1983). In fact, it is 
not realistic to focus on preservation of static ecosystems for preservation of 

J Non-equilibrium 

5 and irregularities of all sorts are not 
aberrations, but integral parts of nature (Solbrig 1991). Both chance and 
natural selection produce the steady coming and going of species through 
a pool. Speciation, immigration, and extinction function continuously. Thus, 
the size of a species pool (species richness) is indeterminate and unpredict- 
able. The theory of species equilibrium, namely, that the steady-state num- 
ber of species, found on an island or isolated patch of habitat due to a 

cause It denies the reality c 
Heterogeneity and No 

The basic arguments of the currently dominant energy liypothesis are: (1) 
species richness is a measure of available energy, and (2) the product of a balance 
between immigration and extinction which causes richness to approach its theo- 
retical maximum over time (MacArthur and Wilson 1967). Therefore, this hy- 
pothesis is that species richness in similar environments is the result of inevi- 
table convergence. In general, tree species richness supports this and increases in 
direct relation to precipitation or AET, suggesting that a positive relationship 
exists between diversity and productivity of the habitat (Latham and Ricklefs 
1992). But this is not always true. There are many examples of species richness 
decline at high habitat productivity levels (Ricklefs 1987, Latham and Ricklefs 
1992). The total number and species of plants within a community may reflect 
total productivity, but species richness does not. Different species have dif- 
ferent individual size, population and distribution patterns and thus have 
different energy needs and different ecological roles in an ecosystem. Some 
species, known as keystone species (Wilson 1992), affect the survival and 
abundance of many other species in the community in which they live. In 
contrast, the presence of some species may be caused by, and be largely 
dependent on, the existence of other species. 

Clearly, species is not an energy or ecological unit. Theoretically, every spe- 
cies has a unique niche because every species has at least one physical or behav- 
ioral characteristic that defines it from other species (Solbrig 1991) even though 
the current measures of environmental parameters are too crude to distinguish 
all differences among species. The ecological inequality of species indicates that 
the number of species present is not a function of the physical environment. On 
the other hand, all regions exhibit heterogeneity and patchness and no environ- 
ment is completely homogeneous. This variability and patchness in the envi- 
ronment provides a foundation for the coexistence of species. Consequently, 
competition seems less important as a determinant of species richness 
throughout a large scale biogeographic region than on a local site. How- 
ever, species richness does not always tend toward its possible maximum 
largely due to the effect of chance. There is reason to assume that nowhere 
in the world have resources been fully utilized by plants. A measure of 
energy availability cannot even allow prediction of the maximum number 
of species in a region because plants have the ability to expand local envi- 
ronmental constraints during colonization. If species saturated biological 
communities exist within limits set by local conditions, new species could 
not join the community without the compensating disappearance of others 

Li and Adair, Species pools 297 

(Ricklefs 1987). But many successful introductions of exotic species can- 
not be shown to have caused an apparent loss of native species. In subtropi- 
cal eastern Asia, for example, where tree species richness is about four times 
that of subtropical eastern North America, about 50 species of trees and 
woody vines introduced from eastern North America have successfully colo- 
nized without the compensating disappearance of the native species. Ameri- 
can Robinia pseudoacacia L., Campsis radkans Seem., Sabina virginiana (L.) 
Antoine, Pinus elliottii Engelm., P. taeda L., P. rigida Mill., Magnolia gran- 
diflora L., Gary a illinoenm (Wangenh.) Koch, and Liriodendron tulipifera L., 
all now grow well in China and do even better locally than their eastern 
Asian species pairs. Of them, Robina pseudoacacia and Campsis radicans have 
escaped from plantations and gardens and became naturalized species in 
China (He and Gu 1990). In these cases, competition between exotic and 
native species has lead to increased diversity rather than extinction and loss 
of species in the area. Thus, a climate or energy hypothesis alone cannot 
explain this fact of additional species richness because both neglect histori- 
cal plant evolution and migration. 

Previous studies have usually concentrated on a small taxonomic group 
of organisms and thus failed to provide a general theory for the existence of 
species pools. In short, what is present on a given site at a given time, or 
interval of time, is a product of chance. The natural processes flowing in a 
river of information are relatively deterministic, but the residents on site at 
a given time are only those who then happen to reside there. 

We are grateful to Barney L. Lipscomb of Botanical Research Institute of 
Texas, Hebe R. Mace, Darrel L. McDonald, Elray S. Nixon, Hershel C. Reeves, 
and Kenneth G. Watterston of Stephen F. Austin State University, and an anony- 
mous reviewer for reading the draft and providing valuable comments and 

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SwiNK, Floyd and Gerould Wilhelm. 1993. Plants of the Chicago Re- 
gion. Fourth Edition. (ISSN 1-883362-01-6, hbk). Indiana Academy 
of Science, 1 102 North Butler Avenue, Indianapolis, IN 46219-2918. 
$28.00 (Academy members); $35.00 (non-members) (+ postage: $2.50 
domestic, $5.00 foreign). 921 pp. 
The "Chicago Area," lying within a 75-mile radius of the Windy City's center, includes parts 
of Illinois, Indiana, Michigan, and Wisconsin. The first (1969) and second (1974) editions of 
this work were by Swink alone; the third (1979) added Wilhelm and well-done keys. This 
fourth edition, with 226 more taxa than its predecessor, is the most splendid of them all. Even 
though Chicagoland is urbanized and depressingly industrialized, it still has many "natural" 
areas and a flora of 2530 taxa, about 35% of them introduced. The arrangements of families, 
genera, and species is strictly alphabetical, as in an ordinary dictionary. Look, then, for the key 
to genera of Cyperaceae under the C's; the key to, and species entries for, Sarpus under the S's. 
The species accounts include not only habitat/historical/phenological/taxonomic/nomenclatural 
data, but also listings, often lengthy, of associated species. The glossary is well illustrated; the 



Laboratory of Systematics and Evolutionary Botany 

Institute of Botany, Chinese Academy of Sciences 



Laboratory ofPhylogenetic Botany 
Faculty of Science, Chiba University 
1-55 Yayoi-cho, Chiba 260, JAPAN 


Hangzhou Botanical Garden, Taoyuanling 

Hangzhou, Zhejiang 510015 


identifications. Based on taxonomic and karyological investigations and crossing tests, 20 
cies are recognized of which four are regarded as hybrids: Lycoris xhaywardii, L. Xa/biflor, 
xhoudyshelii, and L. xrosea. Lycoris aurea var. angustitepala is proposed as new. Keys, descripti 
synonymies, typifications, and karyotypes are included. 

The genus Lycrms Herb, consists of 20 species distributed in warm temperate 
3 subtropical zones of East Asia from southwestern China to Japan and south- 
rn Korea, with a few extending to northern Indochina and Nepal. The species 
re easy to hybridize with each other, and a number of the presently recognized 
ixa are certainly of hybrid origin. 

^resent address: Department of Environmental and Resources Biology, School of Life Sciences, 

Introduction and cultivation ofLycoris species began in the 18th century, first 
to England and then to the United States. To date, most of the species have 
proven successfial in cultivation. They are summer- and autumn-blooming plants 
of great beauty and common in the gardens of China, Japan, and the United 
States. In comparison with other well-known bulb flowers, such as narcissi and 
lilies, Lycoris has its own characteristics and merits. Lycoris comes into flower at a 
time when few other bulbous plants are active. The flowers are characterized by 
their pastel and plentiful colors as well as by beautiful and varied shapes. The 
plants are tolerant to drought, waterlogging, and pests, growing vigorously 
even in barren land, and thus show good prospects in horticulture. 

The genus Lycoris was founded by Herbert in 1821, and L. aurea (L'Her.) 
Herb, was assigned to be the type. In the mid 19th to the early 20th century, 
nine new species were published by various European authors: L. afrkana (Lam.) 
M.J. Roem. (1847), L. straminea Lindley (1848), L. sewerzowtt Kegel (1868), L. 
squamigera and L. sanguinea Maximowicz ( 1 88 5), L. terraccianii Dammann (1889), 
L. sprengeri Comes ex Baker (1902), L. incarnata Comes ex Sprengel (1906), and 
L. argentea Worsley (1928). In the first half of the 20th century, three new spe- 
cies from Japan were added: L. albiflora Koidzumi (1924), L. koreana Nakai 
( 1 930), and L. kiusiana Makino (1 948). After that, the American botanist Hamil- 
ton P. Traub focused his research on the tribe Amarylleae, Amaryllidaceae, and 
published 10 new species (two with iVIoldenke) oi Lycoris based mainly on mate- 
rials introduced from China and Japan and cultivated in American gardens: L. 
m&z (1949); L. haywardu, L. caldwellii, L. houdyshelii, and L. Xw^oo^/V (1957); L. 
chimnsh and L. e/j/^^ (1958), L. Y^lajolla (1963), L. xjacksomana (1964); and L. 
josephinae 0-965). Two keys to the species of the genus were presented in his two 
issues (Traub 1957, 1958). Two American horticulturists, Hayward and Caldwell, 
had done much of work on introduction and cultivation o^Lycoris species. Hay- 
ward (1957) published a new species: L. trauhti. After an interval of about 1 5 
years, the Chinese botanists Hsu & Fan (1974) and Xu et al. (1982) published 
four new species of Lycoris from China: L. iongituba, L. anhuiensis, L. guangxiensis, 
and L. shaanxiensis (1982). Their revision of the Chinese Lycoris, in which 1 5 
species were included , was published in Flora Reipublkae Populans Sinicae (1985). 
Recently, M. Kim and S. Lee (1991) published a new species, L.flavescens, m 
their study of Korean Lycoris. To date then, 29 species have been published. 
However, some of these have been reduced to synonyms, some to infraspecific 
rank, and still some are cultivars or hybrids. Twenty species are included in the 
present paper, though the identity of a few of them is still equivocal and de- 
serves further study. A number of species published in the earlier period are 
without indications of type specimens or even precise type localities; many type 
specimens are not available to us. Besides, the classification of the genus Lycoris 
is largely based on morphology and color of flowers; leaf morphology is not of 
much value in identifications. Furthermore, Lycoris species are easy to hybridize 

Hsu, Synopsis of Lycoris 303 

with each other, and hybrids of diverse morphological features occur frequently 
in nature and in cultivation. All this has caused great difficulties in our research 
work. In many cases, karyological investigation furnishes a taxonomic identifi- 
cation with solid data. 

Lycoris belongs to the subtribe Lycorinae of the niheAmarylleae of subfamily 
Amaryllidoideae. Ungernia Bunge, the only other genus of the subtribe, differs 
from Lycoris mainly in its stamens inserted in two rows instead of one row near 
the throat of the tepaltube, and in its basic chromosome number, x= 1 1 . Ungernia 
has six species with a distribution confined to Iran and central Asia. 

Lycoris was divided by Traub and Moldenke ( 1 949) into two subgenera based 
largely on flower form and length of stamens. Species of subgen. Symmanthus 
Traub & Moldenke have wide tepals that are erect or recurved only at the tips, 
with the distal part more like a trumpet lily. In this subgenus, some species have 
slightly wavy-margined tepals, but the majority do not. Fragrance is found only 
among the funnel-form species. Species of subgen. Lycoris have spiderlily-form 
flowers with substantially reflexed tepals, and the stamens and style usually 
long-exserted. The tepals are almost invariably crisped, wavy, or undulate. The 
flowers are not fragrant. 

Phenologically, the species oiLycoris may be divided into two types. Those 
species from areas with colder winters make little or no leaf growth until Janu- 
ary or later. Those species from areas with warmer winters make leaf growth in 
the autumn and remain in active growth all winter. In both cases, leaf growth 
continues actively until late spring, whereupon the leaves die back. The plants 

Cytological studies on Lycoris were initiated in the late 1920s. To date, chro- 

most of the species. There are three major chromosome types: (1) M (metacen- 
L (acrocentric chromosomes); and (3) T (telocentric chro- 
1 the species with 2n = 22, all rod chromosomes are As; in the spe- 
T 2-16, all rod chromosomes are Ts. The genus has a series of basic 
? numbers — 6, 7, 8 and 1 1 — but the total number of arms of a 
complement of any species is always multiples of 1 1 . It seems that 
1 1 is the primitive basic chromosome number (Liu & Hsu 1 989). However, it is 
still unresolved whether a successive decrease in chromosome numbers as a re- 
sult of Robertsonian fiasion or a gradual increase in chromosome numbers brought 
about by centric fission has been the essential mechanism for karyotype evolu- 
tion and speciation in the genus. Hsu and Liu (1987) proposed a new "fusion- 
fission synthetic theory" for explaining the karyotype evolution and speciation 
in Lycoris. The theory embodies three central ideas: (1) the basic chromosome 
number oiLycoris is x= 1 1 , and 2n=22 is the primitive karyotype; (2) both fusion 
and fission have taken place, at different times during the process of karyotype 
evolution; and (3) duplications, translocations, and other ch 

tions must have occurred in the process of fusion. The meiotic behavior of the 
diploid hybrid L. aff. alhiflora gives evidence for the asstimption that Robertsonian 
changes have occurred in the process of karyotype evolution (Liu & Xu 1990). 
Kurita (1988b), however, insisted that the M type chromosome is not a simple 
product of the fusion of two A type chromosomes. Based on karyological and 
karyogeographical evidences, together with C-banding patterns and DNA con- 
tents of certain species, he is inclined to the fission theory. 

Much work on hybridization and breeding was accomplished mamly by 
horticulturists from the United States, Japan, and China in the past 50 years. 
In the early 1940s, Wood (cf. Adams 1976) crossed L. radiata with L. traubii, 
creating the hybrid L. Xwoodii Traub. Creech (1952) reported the pollination 
of L. radiata by L. aurea; the resulting hybrid had 2n=l9. Takemura (1961, 
I962a,b) made a series of artificial crosses between L. sprengeri, L. straminea 
(probably not true L. straminea), L. radiata, L. radiata \z.r. piimila, L. sanguinea, 
and L. aurea, and investigated morphology and cytology of the hybrids. Caldwell 
(1972, 1981) made many crosses beginning in 1954. One beautiful Caldwell 
hybrid is L.jacksoniana (Traub 1964), the parental species of which are L. sprengeri 
and L. radiata. Koyama ( 1 962) observed meiotic behavior of that hybrid. Adams 
(1976) made numerous crosses, mostly repeating those that had been done 
before. Lin et al. (1990, and unpub.) have been successful in bringing forth 
new hybrids and have shown that both L. rosea and L. haywardii are hybrid 
segregates of L. radiata var. pumila 9 x L. sprengeri S , and that L. squamigera is 
the hybrid of L. chtnensts 9 x L. sprengeri $ . One new hybrid resulting from L. 
sprengeri 9 X L. chinensis S , which they have named L. Xelegans, ined. re- 
sembles L. albiflora in general appearance, but its karyotype is 2?z= 3M+16A= 1 9. 
Diploid species oiLycoris are easy to hybridize with each other, irrespective of 
their conspicuous morphological and ecological differences. The fertility of these 
hybrids is high (Lin et al. 1990). The properties of hybrids can be maintained 
by means of vegetative propagation. This is probably why hybrids often occur 
in nature as well as in cultivation. On the contrary, hybridizations between the 
triploid lycorises are generally unsuccessful (Lin et al. 1990). Results obtained 
from an extensive interspecific crossing program involving reciprocal combi- 
nations of many species by Xu et al. (1986) have shown that both pollen viabil- 
ity and seed-set after crossing are correlated with ploidy level. 

Only a few palynological studies oiLycons have been carried out. Kurita 
(1985) found that there was a latitudinal topocline of variation in pollen 
ornamentation as well as in the amount of gemmate protrasion on the foot 
layer of pollen grains within Lycoris sanguinea var. sanguinea in Japan. As a 
result of a palynological study of five species oi Lycoris from southern Ko- 
rea, Lee and Kim (1987) conclued that pollen size, muri thickness, lumina 
size, and lumina number per 100 pm^ of these species were fundamentally 
in direct proportion to number and size of chromosomes. 

Hsu, Synopsis of Lycoris 305 

Because many investigations of karyology, morphological variation, distri- 
bution patterns, and breeding ofLycons species have been carried out in recent 
years, Traub and Moldenke's revision (1949) requires modification. It is hoped 
that the present revision of the genus will contribute to this objective. 

LYCORIS Herb., Bot. Mag. 47:t.2 113. 1821. Bench. &Hook.f., Gen. PI. 3:727. 

1883; Maxim., Bot. Jahrb. 6:73. 1885; Baker, Handb. Amaryll. 39. 1888; Spreng., 

Bull. Soc. Tosc. Ortic. 8:323. 1888; Traub & Moldenke, Amaryllidac: Tribe Amaryll. 

165. 1949;Traub, PI. Life 13:42. 1957 & I.e. l4:42. 1958; Ohwi, Fl. Jap. 383. 1978; 

Hsuetal.,Fl.Reipub.Pop.Sm. 16:16. 1985;Kim&Lee,KoreanJ.Pl.Taxon. 21:3. 

Bulbous perennial herbs. Leaves after or before the flowers, strap-shaped or 
linear, basal, dying away before flowers develop. Flowers showy, red, yellow, 
peach, lilac and blue, or white (intermediates and pastel tones of the colors are 
also present in some species and hybrids), subtended by 2 spathe- valves, in an 
umbel on a solid scape; perigone 6-lobed, fiinnel-form and regular or spiderlily- 
form and irregular; tepaltube short but expanded at top, throat bearing scales; 
tepals clawed, reflexed at the tips or not; stamens 6, inserted near throat, declinate, 
exserted or not; style long, with a very small capitate stigma, ovary inferior, 3- 
loculed, ovules few in each locule, placentation axile. Fruit a loculicidal capsule; 
seeds few, round, smooth, black. 

There are about 20 species native to China, Japan, and southern Korea, with 
a few extending southwestward to northern Indochina and Nepal. 
1 . Flowers funnel-form, regular; repals not crisped-margined or only minutely 
wavy at base, erect or slightly recurved at the tips; stamens included or slightly 
exceeding tepals (Subgen. Symmanthm Traub & Moldenke). 

4. Flowers produced in Aug.-Sep., vivid rose in the throat, ink-blue at the 
tips of tepals, otherwise purplish rose (China) 1 

4. Flowers produced in Jul -Aug , bluish-mauve with a deeper mauve 
keel (Upper Burma) i 

let with a deeper keel and changing to ink-bluc at tips of tepals (Chma, 
Japan, only cult ) 3 L 

2. Leaves usually appearing in earlv spring flowers never tinged vvitli blue 

1-2.5 cm long; flowers flesh-colored or light rose. 

6. Flowers light purplish pink; tepals 6.3-7 cm long (China, Japan, 

Korea) 4. Z 

6. Flowers nor light purplish pink; tepals 5-7 cm long. 

7. Flowers white, changing to flesh-colored or light rose; tepals with a 

8. Tepals slightly recurved; stamens shorter than or subequalling 

tepals; flowers in lateJul.-Aug. (Japan) 6a. L. sanguineavzt. sanguima 

9. Flowers 7-8 cm long (Japan) 6b. L. sangumea^^,. ktustana 

9. Flowers 5-6 cm long (Korea Japan) 6c. L. sangmnea var. koream 

5. Leaves 1.5-2.5 cm broad, with a distinct whitish stripe; tepaltube 2-6 cm 

10. Tepaltube 2-3.5 cm long; flowers yellow (Chma) l.L.anhinensn 

10. Tepaltube 4.2-6 cm long; flowers yellow or white to peach-colored. 
11. Flowers white with light reddish stripes or peach-colored in bud 

and openi ng pi nkish (Ch 1 na) 8a. L. longiwba var. longituha 

11. Flowers yellow (China) Sh.L. longituha y^r.flava 

. Flowers spiderlily-form or nearly so, distinctly irregular; tepals strongly or me- 
dium crisped-margined, strongly recurved at the tips (Subgen. Lycoris). 
12. Leaves produced in early spring; stamens shorter than to slightly exceeding 

13. Leaves without a distinct whitish stripe; stamens shorter than tepals. 
14. Flowers m bud peach-colored, opening to creamy yellow, changing 

14. Flowers white with a few pinkish stripes inside and a reddish keel 
underside of each tepal (China) 10. L. shaanxiensis 

13. Leaves with a distinct whitish stripe; stamens subequalling or slightly 

15. Leaves narrow strap-shaped, dark green, 24-29 cm long, 1-1 .2 cm 

broad ; tepals with redd ish bands (China) ILL. guangx'umh 

. Leaves produced in autumn; stamens usually far exceeding tepals. 
16. Leaves more than 1 2 mm broad; flowers various colored, but never n 

19. Tepals 7-12 mm broad. 

20. Tepals upright (Upper Burma) \lh.L.aima\ 

19. Tepals 4-8 mm broad (China) Vhz.L. aimav^^. an 

more or less horizontally spreading, rich orange-yellow; tepals 
with a deeper orange-yellow band underside, narrow oblanceolate, 
13-16 mm broad; spathe-valves ovate, 3.5 cm long; pedicels 

Flowers pale straw-colored, creamy white to white, or salmon-colored. 

23. Flowei 

Hsu, Synopsis of Lycoris 

yellow band in the center; tepalcube ca. 2 cm long (Ja 

pan, Korea, Chma) 15. 

23. Flowers creamy white, changing to whitish; tepals wit 
a greenish band underside; tepaltube ca. 8.5 mm lon^ 

(China) 16.1 

22. Flowers pale straw-colored; tepals with a pink band and 
few scattered red dots in the upper surface, changing t 
white in full blossom; tepaltube 4-5.5 mm long (China 

21. Flowers salmon-colored, finally fading to a flesh-color; tepals 
with a deep purplish band tinged with creamy and yellow along 

the center; tepaltube 12-13 mm long (Japan) 18. L. ehiae 

16. Leaves less than 9 mm broad; flowers bright red or rose-colored. 

24. Flowers bright red or sometimes with white-margined tepals; tepals 
distinctly crisped-margined and strongly reflexed; stamens 2-2.5 
times the length of tepals. 
25. Flowers bearing no seeds (China, Japan, Korea, Nepal) 

25. Flowers bearing seeds (China) 1 9b L. radiata var, pumila 

crisped-margined or only ruffled at the bast, slightly reflexed; sta- 
mens a little longer than tepals. 

26. Flowers pale red or becoming whitish, with tepals 1.5-3.2 cm 

long and 2-5 mm broad (Japan) 1 9c . L. radiata var. kazukoana 

26. Flowers rose-colored, with tepals 4-6 cm long and 7-8 mm 

broad (China) 20. L. Y^rosea 

1. Lycoris sprengeri Comes ex Baker, Gard. Chron. ser. 3, 32:469- 1902. Traub 
&Moldenke,Amaryllidac.: Tribe Amaryll. 170. 1949; Traub, PI. Life 13:43. 1957, m 
clavis; Hsu et al., Fl. Reipub. Pop. Sin. 16(1):24. 1985. Type: HUBEI PROVINCE: 
mountains near Xiangyang, not indicated. Sprenger (1906, cf Traub & Moldenke 1949) 
stated this species was sent to him by his collector from China about 1900. They (1949) 
remarked that Comes' reference, which had never been found, was probably a specimen 

Leaves ensiform-linear, ca. 30 cm long and 1 cm broad, dark green. Perigone 
vivid rose in ttie throat, otherwise purphsh rose with ink-blue tips on the tepals. 
Tepaltube 1.2-2.3 cm long; tepals oblanceolate, 4.5-7 cm long, 1-1.7 cm wide, 
not ruffled -margined, recurved. Stamens somewhat shorter than tepals. Style 
about as long as or slightly exceeding tepals. 

Phenology: leaves appearing in early spring; scape produced in August to 

Karyotype: 2«:.22A:=22 (Kurita 1987a; Liu & Hsu 1989). 

Distribution: endemic to China (Anhui, Hubei, Jiangsu, Zhejiang). In bam- 
boo groves and on moist slopes in sparse woods; ca. 100 m. 

Specimens examined: CHINA. Anhui: Chuxian, Langya Shan, East China Bot. Station 5114 
(JSBI). Jiangsu: Yixing, W.Z.Fang 297 (PE); Y.L.Keng 2539 GSBI); yy^ngnmg, J.S.Yue 
0561(JSBI); Shanghai, TianmaShan, G.J. Fans.n. (PE); same locality, Heng Shan, G.J.Fans.n. 

(PE); S.SSu 253 (JSBl); Nanjing Umv. Exped. 255 (SZ); same locality, Songjiang, G.J. Fan s.n. 
(PE); without precise locality, W.C. Chow 819 (Pl^S). Zhejiang: Xiaoshan, Y.L.Keng 11 U (PUS); 
Hangzhou, Hangzhou Bot. Gard. cult., Z.Z. Yu 004, 012, 013 (HZBG); Zhoushan,/ Z.L/k 
008 (HZBG). 

This species shows much variation in breadth of tepals and in lengths of 
tepaltube and stamens. The plants coming from Zhoushan, Zhejiang, show 
some differences in length of stamens, some with stamens shorter or subequalling 
the tepals, some with longer and exserted stamens. 

2. Lycoris argentea Worsley, Gard. Chron. sen 3, 84:169, fig. 72. 1928. Grey, 

Royal Horticultural Society, England, in October 1904 by C. Judes from Upper Burma. 

Leaves bluish green. Perigone bluish mauve with a silvery sheen and some 
sparkles and a deeper mauve keel. Tepaltube very short; tepals oblong, neither 
undulate-margined nor recurved. Stamens and style subequalling tepals. 

Phenology: leaves appearing in autumn; scape produced in July to August. 

Karyotype: no reports. 

Distribution: Upper Burma. 

The above morphological description is based on Worsley 's brief description 
of the type. No other detailed information is available. According to Worsley 
(1928), this species differs from L. squamigera Maxim, in having bluish-mauve 
tepals with a deeper mauve keel, and in the stamens and style being about equal 
to the tepals. We have not seen any specimens of this species at Kew. It has 
probably a more close affinity to L. xhaywardii than to L. squamtgera, at least in 
morphology. At present, the identity of this species, be quite equivocal, de- 
serves further study. 

3. Lycoris xhaywardiiTraub(prosp.), PI. Life 13:44, fig. 16. 1957. Caldwell, 
PI. Life 13:53. 1957. Type: UNITED STATES. Tennessee: Nashville (cult.), S. Caldwell 
554 (holotype: TRA). Paratype: FLORIDA: Winter Park, W. Hayward291 (TRA). 
According to Caldwell (1957), the type was from Japan. 

Leaves up to 48 cm long, 7-1 1 mm broad, deep green, glaucescent. Perigone 
reddish violet, a little lighter than in L. sprengeri, tepals with a deeper keel and 
changing to ink-blue at the apex. Tepaltube 1.1-1.3 cm long; tepals oblan- 
ceolate, 4.4-5.5 cm long, 1-1 . 1 cm broad, not crisped-margined. Stamens some- 
what shorter than tepals. Style moderately exserted. 

Phenology: leaves appearing in autumn; scape produced in July to August. 

K^O/()^)//?e; 2«=22A=22 (Hsu etal. 1981; Liu& Hsu 1989). 

Distribution: known only in cultivation. 

Specimens examined: CHINA. Zhejiang: Hangzhou, Hangzhou Bot. Gard., cult.,7.Z.L/« 
002, 003, Z.Z. Yu 026 (HZBG). The specimens cited differ from the type in having longer 
stamens exceeding the tepals. 

Hsu, Synopsis of Lycoris 309 

This species is most closely allied with L. sprengeri Comes ex Baker and differs 
from that species mainly in having smaller flowers with a paler bluish purple 
color and an earlier blooming time. 

Hybridizing tests accomplished by Lin et al. (unpub.) have shown that L. 
xhaywardii is a hybrid between L. sprengeri and L. radiata ydx.pumila. The arti- 
ficial hybrids resemble L. xhaywardii very much in external morphology. 
Karyologically, both of the parental species have 2n=22h., also in accord with L. 

4. Lycoris squamigera Maxim., Bot. Jahrb. 6:79- 1885. Baker, Handb.Amaryll. 

Nakai, Fl. Kor. 234. 1 9 1 1 ; Worsley, Gard. Chron. ser. 3, 84:169. 1928; Traub & Mold- 
enke, Amaryllidac: Tribe Amaryll. 173. 1949; Koyama, Baiieya 7:6, fig. 18. 1959; 
Ohwi, Fl. Jap. ed. 2,384. 1978; Hsu et al., Fl. Reipub. Pop. Sin. 16(1):24. 1985; M.Kim 
& S.Lee, Korean. J. PI. Taxon, 21:10. 1991. Type: JAPAN: Maximowicz cited three 

AOT^ry/H/)rf///zHovey ex Baker, Bot. Mag. 123:t.7547. 1897,insyn. 

Leaves 1 .8-2.5 cm broad, bright green. Perigone light purplish pink with a 
gold throat. Tepaltube 1 .6-2.5 cm long; tepals oblanceolate, 6-7 cm long, 1 .2- 
1.8 cm broad, minutely wavy-margined at base, not reflexed. Stamens 
subequalling tepals. Style slightly exserted. 

Phenology: leaves appearing in spring in Japan and Korea, in autumn in China, 
then dying down and regenerating in spring; scape produced in August. 

Karyotype: 2;2=6M+ 10T+ 1 1A=27 (Kurita 1987a; Liu & Hsu 1989). 

Distribution: eastern China (Jiangsu, Shandong, Zhejiang), Japan, and Korea. 
In moist often disturbed places, such as margin of plantations, around dwell- 
ings, and graveyards; to 1200 m. 

Specimens examined: CHINA. Jiangsu: Yuntai Shan, F.X.Uu 10732 (PE); X.R.Changetal. 
19608 (SHMI); Shanghai, cult., P.S.Hsu 486 (FUS, JSBI); C.N.Yan 10391 (FUS). Shandong: 
Qingdao, Lao Shan, H.B.Cm376 (PE). Zhejiang: Shonshan, Y.L.Kengllll (FUS). 

JAPAN. Chiba Pref.: Chiba Univ., cult., S.Kurita 910820 (CBM). Nagano Pref.: 
Kamiminochi-gun, Tokakushi-mura, S.Kurita 870810 (CBM). Tokyo Pref.: Hachioji, 
Kamiange, S.Kurha 860813 (CBM). 

KOREA. Kangwon-do: Kangwon, Mr. Kumgnngsan, T.Uchiyamas.n. (TI). Kyonggi-do: 
Suwon, H.Uekis.n.iTl). 

Based on karyological and morphological studies, Inariyama (1948, 195 1 , 
1952,1953) considered this sterile species a triploid hybrid between L. straminea^ 
and L. sprengeri. Takemura (1961) crossed L. straminea^ and L. sprengeri. The 
hybrids resembled L. squamigera in gross morphology, but they were diploids 

having 2«= 19. Based on cytological investigation, Kurita (1987a) supported 
Inariyama's proposition and was of the opinion that since L. squamigera occurs 
only in human habitations or as an escape in Japan, it was brought to Japan 
from China for ornamental purposes in early time. 

5. Lycoris incarnata Comes ex Spreng., Gartenwelt 10:490, fig. 1 . 1906. Worsley, 

Gard. Chron. set. 3, 84:169. 1928; Traub & Moldenke, Amaryllidac: Tribe Amaryll. 
172. 1949; Traub, PI. Life 13:43. 1957; Hsuet al., Fl. Reipub. Pop. Sin. 16(1):25. 1985. 
Type: no type specimen cited. Sprenger stated that L. incarnata was first described by 

Moldenke 1949). According to Sprenger, this species was collected in Hubei, China, and 
sent to him in Naples about 1901 by a collector. 
Leaves strap-shaped, ca. 50 cm long, ca. 1 .2 cm broad, dark green. Perigone 
white in bud, opening white, changing gradually to flesh-colored or light rose. 
Tepaltube 1 .8-2 cm long; tepals oblanceolate, with a reddish stripe above and a 
deeper keel below, 5-5.6 cm long, 1 .2-1 .4 cm broad, barely undulate-mar- 
gined, only very slightly recurved. Stamens purplish, subequalling or some- 
what shorter than tepals. Style purplish, a little exceeding tepals. 

Phenology: leaves appearing in spring; scape produced in September to 

/C^ryo/y^^e: 2;2=4M+3T+22A+lm = 30 (Kurita 1987a). 

Distribution: endemic to China (Hubei and Yunnan). 

This species is most similar to L. squamigera Maxim, from which it differs 
mainly in the leaves appearing only once in spring, in the size and color of 
flowers, and in the karyotype. 

Kurita (1987a) strongly suggested that L. incarnata must have originated 
from a cross between a gamete having 4M + 3T and another gamete having 
22A+lm, though the parental species could not be decided. 

6. Lycoris sanguinea JVLaxim., Bot. Jahrb. 6:80. 1885 . Baker, Handb. Amaryll. 40. 

1888; Spreng., Bull. Soc. Tosc. Ortic. 8:328. 1888; Worsley, Gard. Chron. ser. 3, 84:169. 
1928;Makino,Illus. Fl.Jap. fig. 2165. 1949; Traub & Moldenke, Amaryllidac: Tribe 
Amaryll. 175. 1949; Koyama, Baileya 7:5, fig. 2a. 1959; Ohwi, Fl.Jap. ed. 2, 384. 

6a. Lycoris sanguinea var. sanguinea, Type: Maximowicz listed two type specimens 
of this species m his type description. They were all from Japan: one near Yokohama 
{Tschonoski fl.), and the other between Kyoto and Maizuru {Doederlein fl., in Engler's 

Leaves strap-shaped, 20-30 cm long, 10-12 mm broad, light green. Peri- 
gone apricot-orange. Tepaltube 1-1.5 cm long; tepals linear-oblong, acute, 6.4- 
7 cm long, ca. 3.8 cm broad, neither crisped-margined nor recurved. Stamens 
subequalling or slightly shorter than tepals. Style exceeding tepals. 

Hsu, Synopsis of Lycoris 311 

Phenology: leaves appearing in spring, dying down and coming up again in 
next spring; scape produced in late July to late August. 

Karyotype: 2« = 22A=22 (Inariyama 1931, 1937, 1951;Koyama 1954, 1962; 
Kurita 1989;Nakamura 1978;Nishiyama 1928; Sato 1942;Takemura 1961; 
Yoshida 1972); 2« = 31A+1M=32 (Kurita 1989). 

Distribution: ^Q-pQ-n (northern Honshu and southwestward, Shikoku, Kyushu). 
In moist places by streams and sparse woods; to 500 m. 

Specimens examined: JAPAN. Chiba Pref.: Chiba, Noromachi, S.Kurtta 870810 (CBM). 
EhimaPref.: Saijyoshi, Mr. Ishizuchi, M.Takahashi 1790 (PE). Kochi Pref.: Takaoka-gun, 
Hayama-mura, H.Ohashi 660203 (TI); Kami-gun, Tosayamada-cho, Ryuga-dou, M.Takahashi 
1 786 (PE). Kyoto Pref.: Yamashiro, Yaze, G .Koidzumi s.n. (TI). NiigataPref.: Awajima, KMon 
47 (TI). SaitamaPref.: Aganomachi, Hanagin, T.Yamazaki s.n. (TI). ShizuokaPref.: Ogasa- 
gun, Kikugawa-cho, S.Kurita 860808 (CBM). 

This species is characterized by its reddish orange or orange-colored flowers 
with stamens nearly equal to or slightly shorter than tepals. But there is a culti- 
var (var. alba Hort. Mill. & Bailey, Stand. Cycl. Hort. 2: 1933- 1939) with white 
flowers and with the leaves appear in March. 

6b. Lycoris sanguinea Maxim, var. kiusiana (Makino) Koyama, Baileya 7:5. 
1959. Ohwi, Fl.Jap. ed. 2, 384. 1978. Type: JAPAN. HizenPref.: Mr. Tarain Kiusiu, 
cultivated in Oidzumi {T. Makims.n.). 

Lycoris kiusiam Mixkmo,Ma.kmoa9:l76. 1948; Komatzuzaki, J.Jap. Bot. 32:62. 1957. 

Differs from L. sanguinea var. sanguinea by the larger-sized flowers (7-9 cm 
long) produced in July with the tepals distinctly recurved, the longer stamens 
distinctly exceeding the tepals, the broader leaves (10-13 mm wide), and the 

Phenology: leaves appearing in February to May; scape produced in early July 
to early August. 

Karyotype: 2« = 22A-22 (Kurita 1988b; Takemura 1965; Yoshida 1972); 
2«=33A&44A (Kurita 1988a). 

Distribution: endemic to Japan (Central Honshu and westward, mainly in 
Kyushu). In shady slopes under forest trees. 

Specimens examined: JAPAN. OoitaPref.: Mt. Kyoyomi-dake, l.Emmotos.n. (TI). Nagasaki 
Pref.: Mt. Tara-dake, H.Haras.n. (TI). Tokushima Pref.: Miyoshi-gun, Nishisofuyama-mura, 
Zentoku, Y.Mofmyama 288 (TI). Wakayama Pref.: Hitaka, Kawakami-mura, Imoo, T.Yamasaki 
s.n. (TI). Kyoto Pref.: Kifline, S.K»ma 860725 (CBM). Tokyo Pref.: Nishkama, Mt. Kagenobu, 
S.Kunta 8407 15 (CBM). 

This variety siiows little difference in character from L. sanguinea var. kiusiam 
from which it differs only by its smaller-sized perigone (5-6 cm long). 

Phenology: leaves appearing in February to May; scape produced in August. 

Karyotype: 2n =22 A^22{^\xnt?i 1988a; Taeetal. 1987;Takemura 1965). 

Distribution: southern Korea and Kyushu and Tsushima Island of Japan. I 
moist places in the mountains; to 600 m. 

7. Lycoris anhuiensis Y.Hsu & G.J.Fan, Aci 

Hsuetal.,Fl.Reipub. Pop.Sin. 16(1):25, pl.7. 
Shan, Y.Xu & G.J.Fan 2234 (holotype: SHMI). 

Leaves strap-shaped, ca. 35 cm long, 2-2.5 cm broad, green with a distinct 
whitish stripe in the center. Perigone yellow. Tepaltube 2-3.5 cm long; tepals 
ca. 6 cm long, 1.3-1 .7 cm broad, slightly ruffle-margined, spreading and some- 
what recurved. Stamens as long as tepals. Style somewhat exserted. 

Phenology: leaves emerging in early spring; scape produced in August. 

Karyotype: 2«=6M+ 10T= 16 (Hsu & Lm 1987). 

Distribution: endemic to China (Jiangsu and Anhui). On stony slopes in the 

Specimens Examined: CHINA. Anhui: Cliuxian, East China Bot. Station 5 163 (PE). 

This species is characterized by its yellow flowers and by its leaves possessing 
a whitish stripe in the center. It resembles L. longituba described below in both 
morphology and karyotype, but differs from that species in having smaller yel- 
low flowers and shorter tepaltube. 

8. Lycoris longituba Y.Hsu & G.J.Fan, Acta Phytotax. Sin. 1 2 :299, pl.6l . 

1974. Hsuetal.,Fl. Reipub.Pop.Sin. 16(1):27. 1985. Type: CHINA, Nanjing, Mt. 
Zijinshan, F.X.Liu 7 97 9 (hokotype:JSBI; isotyph: SHMI). Paratypps: Nanjmg, Mc. 
Baohua 'i>\x^nJ.J.Gong 00869 (JSBI); Jiangning County, Mr. Dachenyishan, M.Z.Choii 
64244 aSBI); Zhenjiang, Zhulinsi, H.Migos.n. (JSBI). 
8a. Lycoris longituba var. longituba 

Leaves lanceolate, ca. 38 cm long and ca. 2.5 cm broad, with a distinct whit- 
ish stripe in the center. Perigone white with light reddish stripes or peach- 
colored in bud and opening pinkish. Tepaltube 4.2-6. 6 cm long; tepals 6-7 cm 
long, 1.5-2.1 cm broad, not ruffle-margined, somewhat recurved. Stamens 
slightly shorter than tepals. Style nearly equal to or slightly exceeding tepals. 
Phenology: leaves appearing in early spring; scape produced in July to August. 
Karyotype: 2n=6M+ 10T= 16 (Liu & Hsu 1989). 
Distribution: endemic to China (Jiangsu and Anhui). 

Hsu, Synopsis of Lycoris 3 13 

Specimens examined: CHINA. Jiangsu: Zhenjiang, East China Bot. Station 2997 (PE); 
Nanjing, F.X.Liu 73 19 (PE); same locality, Sun Yat Sen Mem. Bot. Gatd. cult., Z.Z. Yu 024, 
Z.G.Mao 10501 (HZBG). 

This distinctive species is characterized by its regular, large, white (or with 
reddish stripes) to pinkish flowers with long tepaltube (the longest in the ge- 
nus). But the species shows much variation in the shape and color of flowers 
correlated with karyotype variation. A form with rather thick and greenish pale 
yellow tepals possesses 2w=7M+8T=15; another form characterized by having 
shorter tepals possesses 2?z=7M+6T+2A=15. Since the short arm of this A type 
chromosome is revealed to be heterochromatic, it may be derived from an inver- 
sion of the Tty 

8a. Lycoris longituba Y.Hsu & G.J.Fan var. flava Y.Hsu Sc X.L.Huang, Acta 
Phytotax. Sin. 20:198. 1982, and Fl. Reipub. Pop. Sm. l6(l):27. 1985. 
Type: CHINA. Jiangsu: Jiangnmg, Mt. Langshan, Z.M.Zhou 64246 (holotype: SHMI). 

Flowers pale yellow. 

Phenology: same as var. longituba. 

Karyotype: 2n=6U+ 10T= l6 (Kurita et al. unpub.). 

Distribution: endemic to China (Jiangsu). On slopes of hills of low elevation. 

In Caldwell's (1 979) reference, Fig. 1 1 (unidentified) No. 289 (lefi:) undoubt- 
edly belongs here. The plant was said to have been imported to Japan from 
China in the 1940s or even earlier. Caldwell got this plant from a Japanese 
hobby gardener, Dr. Shuichi Hirao of Kanagawa. 

9. Lycoris caldwellii Traub, PI. Life 13:46, pl.4. 1957. Caldwell, PI. Life 13:55. 
1957; Hsu et al., Fl. Reipub. Pop. Sm. 16(1):24. 1985. Type: UNITED STATES. Ten- 
nessee: Nashville (cult.), S.Caldwell 552 (holotype: TRA); same locality, S.Caldwell 222 

to the United States from a shipment out of Shanghai, China, before 1949. 

Leaves without a whitish stripe in the center. Flowers between the spiderlily- 
form and funnel-form, in bud peach-colored, opening to pale yellow, changing 
gradually to creamy white with age. Tepaltube 2-2.2 cm long; tepals oblan- 
ceolate, 7-7.5 cm long, slightly undulate margined, recurved, 1 .2-1 .4 cm broad. 
Stamens shorter than tepals. Style subequalling tepals. 

Phenology: leaves appearing in early spring; scape produced in late August to 

K^r)/o?)//^e; 2;7=6M+ 10T+ 1 1A=27 (Bose 1957; Liu & Hsu 1989). 

Distribution: endemic to southeastern China (Jiangsu, Zhejiang and Jiangxi). 
Specimens examined: CHINA. Zhejiang: Hangzhou, Hangzhou Bot. Gard. cult.,J.Z. Lin 
07 7 (HZBG). 

The leaves of this species lack a whitish stripe or band in the center, which is 
a distinctive feature of all the species of Subgen. Lycoris. Although the karyotype 

that of L. squa?nigera, the gross morphology i 

10. Lycorisshaanxiensis Y.Hsu &Z.B.HU, Acta Phytotax. Sin. 20:196. 1982. 
Hsuetal., Fl. Reipub. Pop. Sin. 16(1):24. 1985. Type: CHINA. Shaanxi, Mt. Qingling, 

Leaves strap-shaped, ca. 50 cm long and 0.8 cm broad, without a distinct 
whitish stripe in the center. Perigone white, with a few pinkish stripes inside 
and a reddish keel underside of each tepal. Tepaltube ca. 2 cm long; tepals slight 
ruffled-margined, recurved. Stamens shorter than tepals. Style somewhat 

Phenology: leaves appearing in early spring; scape produced in August to 

Karyotype: no reports. 

Distribution: endemic to China (Shaanxi). 

1 1. Lycoris guangxiensis Y.Hsu & G.J.Fan, Acta Phytotax. Sin. 20: 196. 1982. 

H.suecal.,Fl. Reipub. Pop. Sm. I6(l):22,pl.6, fig. 1-2. 1985. Type: CHINA. Guangxi, 
Duan, Y.Xh. & G.J.Fans.n. (holotype: SHMI). The type specimen has not been found in 

Leaves narrow strap-shaped, 24-29 cm long, 1-1 .2 cm broad, dark green 
with a distinct whitish stripe in the center. Perigone yellow, with reddish bands. 
Tepaltube 1.5—2 cm long; tepals obovate-oblanceolate to oblanceolate, ca. 7 cm 
long and 1 . 5 cm broad, narrowing to a claw at base, slightly ruffled-margined, 
recurved. Stamens subequalling tepals. Style exserted. 

Phenology: leaves emerging in early spring; scape produced in July to August. 

Karyotype: no reports. 

Distribution: endemic to China (Guangxi). 

No specimens of this species are available to the present authors. According 
to the type description, this species is most closely related to L. chinensis Traub 
but differs from that species in its perigone having reddish bands inside, and in 
its narrower, dark green leaves. 

12. Lycoris chinensis Traub, PI. Life 14:44. 1958. Hsuetal, Fi. Reipub. Pop. Sin. 

16(1):22, pi. 5, fig. 4. 1985; M.Kim &S.Lee, Korean. J. Pi. Taxon. 21:126, fig. 5-6. 
1987. Type: UNITED STATES. California: La Jolla, cultivated, H.P.Trauh 583 (holo- 
type: TRA). According to Traub's remark, the type plant was grown from a bulb 
{P.I. 162443) furnished by J.L.Creech originally obtained from Sun Yat Sen Memorial 
Garden in Nanjing, Chma, jn 1948. 

. 35 cm long, ca. 2 cm broad, green 

Hsu , Synopsis of Lycoris 3 1 5 

yellowish midrib underside of each repal. Tepalrube 1.5-2.5 cm long; repals 
oblanceolate, 5.5-7.7 cm long, 7-13 mm broad, distinctly crisped-margined 
and strongly recurved. Stamens subequalling or somewhat exceeding tepals, 
filament yellowish. Style rose-colored in the upper part. 

Phenology: leaves appearing in early spring; scape produced in late July to 
August. Fruits in September. 

Karyotype: 2n=6U+lQ)i: =16 {Bose 1960;Chen&Li 1985;Lm&Hsu 1989). 

Distribution: China (Henan, Shaanxi, Sichuan, Zhejiang, and Jiangsu) and 
southern Korea. On moist slopes in the mountains; ca. 750 m. 

Specimens examined: CHINA. Jiangsu: Jurong, Baohua Shm Jiangsu Natural Pi. Sources 
Exped. 5426 (HZBG); Yixing, Y.L.Keng2362 (PE). Sichuan: way from Daheba to Sanquan, 
J.H.Xiong & Z.L.Zhou 92976 (SZ), Zhejiang: Ningbo, Y.L.Keng 1117 (FUS); Tianmu Shan, 
Y.Y.Ho 24601, 24687, 25259, 29332, (HZBG); Hangzhou, S.Y.Chang 858, 7327 (HZBG); 
same locaHty, Hangzhou Bot. Card., cuk., Z.Z. Yu 003,006, 007, 01 1 , 015. 01 7 (HZBG). 

This species resembles L. aurea and L. traubii in general appearance, but dif- 
fers remarkably from them in its foliage appearing in early spring instead of in 
autumn and winter, in the flowers having a chrome yellow color, and in the 
stamens subequalling or slightly surpassing the tepals. Both L. aurea and L. 
traubii bloom in September to October, later than that of L. chinenm. 

The Korean plants, which Tae et al. (1987) have mistaken to be "L. aurea" 
are morphologically very similar to typical L. chinenm ., and the chromosome 
number is also 2n= 16, but the karyotype is slightly different (2«=6M+2SM+8T). 
These SM type chromosome may have originated from an inversion of a T type 
chromosome. So the Korean taxon may be derived from the typical Chinese L. 
chinensis. Bose (1966) also reported a SM type chromosome of the same sort in a 
bulb of L. chinensis, but he did not mention the source of material that he exam- 
ined. A cultivated form called "L. sperryi" has also 2?z = 6M+ 10T= I6. 

13. Lycoris aurea (L'Her.) Herb., Bot. Mag. 47:t.2113. 1821. Hance,J. Bot. 
12:262. 1874;Franch. &Sav.,Enum.Pl.Jap. 2:44. 1879; Maxim., Bot. Jahrb. 6:79- 
1885; Baker, Handb. Amaryll. 40. 1888; Forest, Gard. Chron. Ser.3, 47:12, fig. 15. 
1910;Diels,NotesBot.Gard.Edmburgh6:192.1912;Hayward, PI. Life 13:41. 1957; 
Koyama, Baileya 7:5. 1959; Icon. Cormophyt. Sin. 5:549, fig. 7928. 1976; Hsu et al., 
Fi. Reipub. Pop. Sin. l6(l):20,pl.5, fig. 1-3. 1985. Type: not cited. According to Alton's 
(181 1) reference, this species was introduced ro England from China by J. Fothergill in 
Amaryltis aurea UHei., Sert. Angl. 14, pl.l5. 1788, with descriptions in Hort. Kew 1:419. 
1789; Bot. Mag. 12:t.409. 1798. 

Leaves ensiform, acutish at the apex, up to 76 cm long and to 2.5 cm broad, 
to 76 cm long. Spathe-valves lanceolate, 7-8 cm long. Pedicel 1 5-22 mm long. 

Flowers somewhat upright. Perigone cadmium yellow, with a pale green 
stripe underside of each tepal. Tepaltube 1.2-1.5 cm long, straight; tepals 
narrowly elliptic, 5-6 cm long, 7-12 mm broad, decidedly crisped-mar- 
gined, strongly recurved. Stamens yellowish, surpassing tepals about 1/6- 
1/3- Style exserted, reddish in the upper part. 

Phenology: leaves appearing in autumn in flowering time; scape produced 
in August to September, before leaves. 

Karyotype: 2n=\QM+2T =12 (hose 1958);2«=9M+4T=13(Bose&Flory 1963, 
Flory 1963); 2«=7M+ 1A + 7T= 15 (Kurita 1987a); 2w = 6M+ 10T= 16 (Liu & 
Hsu 1989). 

Distribution: to China and Indochina; in China in southern Henan, Shaanxi, 
and Gansu, southern Jiangsu, Zhejiang, Jiangxi, Guangdong, Guangxi, Hunan, 
western Hubei, Sichuan, Guizhou, and Yunnan. Usually in sheltered moist rocky 
or grassy slopes along streams and at the edges of forests in the mountains; 
(110)500-2250 m. 

Specimens examined: CHINA. Gansu: Wenxian, Q.X.Lt& X.C.Zhao 2420 (PE): Kangxian, 
Z.Y.Zhang 16484 (PE). Guangdong: Guangzhou, cult. Y.Tsiang 13268 (PE); Hongkong, 
Kadoorie farm, brought by Gloria Barretto, S. Y.Hu 11030 (PE). Guangxi: no locahty, Guangxi 
PL Exped. 4034 (PE). Guizhou: Huaxi, cult. Z, Y.Cao 2480 (PE); Jiangkou, southeast side of 
Mt. Fanjing Shan, Smo-Amerkan Guizhou Bot. Exped. 21 7 (PE); Xingyi, Z.S.Zhang & Y.T.Zhang 
6516 (PE); Bijie, Baohuixiang, P.H.Yu 321 (PE); near Panxian, Anshun Exped 1298 (PE); 
Yinjiang, C.B.Jian 30598 (PE). Henan: Jigong Shan, Forest. Depart. Henan Prov. 353 (PE); 
Tongyang, X.Q.Zhang2021 1 (PE). Hubei: Badong, Shennongjia, G.X.Fu& Z.X.Zhang 1 1 70 
(PE); Fangxian, C.L.Cheng 538 (FUS); K.M.Lwu 9065, 9092, 9123 (PE); Enshi, L.Y.Dai & 
Z.H.Qian 620 (PE). Hunan: Yongshan, L.H.Lui 9475 (PE); Anjiang, Agri. School 1466 (PE); 
Nanyue, KLz« 3 76 (PE);Qiangang,Xuefeng Shan, Z.XLi 2721 (PE). Jiangsu: Yixing, W.Z.Fang 
etal. 248 (SZ). Zhejiang: Tianmu Shan, Y.Y.Ho 25259 (PE); T.Tang & W.Y.Hsia 500 (PE); 
Zhejiang PI. Sources Exped 29332 (PE); Hangzhou, S. Y.Chang 1327 (PE); Hangzhou Bot. Card., 
cult. Z.Z. Yu 009 (HZBG); no locality, R.C.Ching 5256 (FUS); 5349 (PE). Sichuan: Wuxi, 
K.L.Chu 1924 (PE); no locality, T.Tang 23625 (PE); Wanyuan, P Y.Li 5584 (PE); Wenchuan, 
S.Y.Chen et aL 5896 (SZ); Yuan, Yaan Exped. 1125 {CDBI); Jieshan, Jmhan Exped. 78-0751 
(CDBI); Jiangjin J/^«^/K Exped. 519 (CDBI); Huidong, Huidong Exped. 341 (CDBI); Huili, 
Tapping Exped. 414 (CDBI); Fengdu, Fengdu Exped. 744 (CDBI); Youyang, Youyang Exped. 506 
(CDBI); Wulong, Wulong Exped. 1302 (CDBI); Pengshm, Pengshui Exped. 1185 (CDBI); Xiushan, 
Xiushan Exped 1177 (CDBI); Leibo, Leiho Med Exped. 967 (CDBI); Xuanhan, Xuanhan Exped. 
1 1 75 (CDBI); Tongjiang, Tongjiang Exped. 756 (CDBI); Lingshui, Lingshui Exped. 965 (CDBI); 
Guanxian, Qingchen Shan, Z.L. Wu 33955 (PE); Bushi, Skhuan Econ. PL Exped. 5803 (PE); way 
from Daheba toYuc[u&n,J.H.Xiong& Z.L.Zhou 92976 (PE); Chengkou, T.L.Dai 102283 (PE); 
Jiange, T.N. Liou ^ C.Wang 293 (PE); Mt. Emei, T.YChow & G.J.Xu 586 (PE); Hechuan, 
Huaging Shan, T.H.Tu 5202 (PE); Huidong, Q.S.Zhao 5 799 (SZ); H.N. Wang 79626 (FUS); 
without precise locality, F.T.Wang 22305 (PE); Ebian, C.W.Yao 3016 (PE). Shaanxi: Ziyang, 
near Liangshuang River, P Y.Li 6382 (PE); Taiba Shan, T.S.Chen 3684 (FUS). Yunnan: Jianchuan, 
Mekong Divide, G.Forrest 23567 (PE); Gongshan, TTYu 23023 (PE); Kunming, T.N.Lwu 
16381 (PE); Shuangbai, H.T.Tsai 54346, 54601 (KUN, PE, SZ); Qiujiang Valley, TTYu 19912 
(PE); Jianchuan, R. C. Ching 23 1 03 (PE); Lunan, B.Y.Qtu 55917 (PE); Yiliang, Northeast Yunnan 

Exped. 944 (PE); Gongshan, Qinghat-Xizang Exped. 9^57 (PE); Jianchuan, R.C.Ching 23103 
(KUN); Qiubei, Kunming BoL Inst. 62-3777 (KUN); Anning, C.Y.Wu231 (KUN); Yiwu, 
SJ.Pei ^9-10108 (KUN). 

Redoute (1822), followed later by Kunth (1850) and Traub and Moldenke 
( 1 949), treated Amaryllis afmana Lam . ( = Lycoris afncana (Lam.) , MJ . Roem . ) as 
a synonym oiA.aurea L'Her. (=L. aurea (L'Her.) Herb.). Recently, some authors 
(e.g., Bailey & Bailey 1976; Everett 1983) have adopted Lycoris afrkana (Lam.) 
M.J. Roem. as the valid name over L. aurea on the basis of the fact that A. 
afncana Lam. (1783) antedates A . ^^re^ LHer. (1788). But it should be noted 
that according to Lamarck's (1783) original description, A. ^/ric^^z^ was culti- 
vated in a garden in Roi and was said to have been introduced from Africa and 
Madagascar, out of the range as so far known of L. aurea. From a morphological 
point of view, these two names may refer to the same entity. But we are not 
making the substitution at the present time pending further studies on type 
material of the taxa involved. 

This subtropical species has a relatively wide distribution extending into 
southeastern Asia, China and Indochina. It is characterized by its long and broad 
ensiform leaves almost acute at the apex, its cadmium yellow flowers with strongly 
crisped and recurved tepals, and its moderately exserted stamens. But some of 
the collections from Sichuan {V. T. Wang 22303 and C. W. Yao 3016) have long 

Five cytoraces (2^2=12, 13, 14, 15, 1 6) under the name of L.^^^r^^ have been 
reported by various authors (Bose 1958; Bose and Flory 1963;Inariyama 1931, 
1932, 1937; Kurita 1987a; Liu & Hsu 1989). But as a result of a reexamination 
of these cytoraces by Kurita (unpub.), only three of them have been confirmed: 
2;2=l4 = 8M+6T,2?z=15 = 7M + 8T,and2;z=l6=6M+10T.Thesecytoracesare 
morphologically distinguishable from one another. The 2«= 1 5 cytorace has as- 
cending tepals and very narrow leaves acute at the apex. The description of L. 
aurea var. surgens Worsley ex Traub & Moldenke as well as the figures of L. 
afncana (Lam.) M.J. Roem. published in Encyclopedia of Horticulture of New York 
Botanical Garden (Everett 1983) match well with this cytorace. Kurita (1987a) 
suggested that this highly sterile cytorace might have originated either from 
the cytorace having 2/2=16 through centromeric fusion between two T type 
chromosomes, or might have been produced by crossing between two fertile 
cytoraces or species having 2n= l4 and 2n^l6. In the 2«= 16 cytorace, the leaves 
are pendulous and are the largest among all cytoraces (60-80 cm long and 4-6 
cm broad) with a blunt apex. This cytorace may be a rheophyte adapted to 
riverside habitats. The size and shape of leaves of the 2«= l4 cytorace are in the 
middle of the above two cytoraces, but the midrib on the under surface of leaves 
is reddish purple, and the central part of each tepal is greenish. Kurita (unpub.) 
suggested that these cytoraces might not be the simple products of Robertsonian 
changes of a single species. 

For a long time, taxonomists have confused this species with L. trauhii Hay- 
ward, which occurs only in Taiwan and southern Japan (Kurita 1980). Many 
specimens identified to be "L. aurea" in Japanese herbaria belong to L. traubii. 
Hay ward (1957) and Kurita (1980) made morphological comparisons of these 
two species. Kurita et al. (unpub.) recently added that the remains of leaf-bases 
is prominent at the base of scapes of L. aurea, whereas in L. traubii there is no 

13b. Lycoris aurea (L'Her.) Herb. var. surgens Worsley ex Traub & Moldenke, 
Amaryllidac: Amaryll. 180. 1949- Type: UPPER BURMA: cultivated at the 
Royal Hortiucltural Society of England,, Oct 1 904 (Worsley 1 928). 

All of the tepals ascending. 

Karyotype: no reports. 

Distribution: Upper Burma. 

No specimens of this variety are available to the present authors. Worsley 
(1928) and Traub and Moldenke (1949) remarked that the foliage is distinct, 
but they gave no further descriptions. This variety is probably in accord with 
the cytorace of L. aurea having 2;? = 1 5 . 

13c. Lycoris aurea (L'Her.) Herb. var. angustitepala Hsu, Kurka, Yu, & Lin, 
A varietate tf//re:^ tepalis angustioribus 4-8 mm latis, stammibus longioribus tepalum 1/3- 

This variety differs from var. aurea mainly in having narrower tepals and 
long-exserted stamens. 

Type: CHINA. Hubei: Fengxian, Guanyindong, 88 m, in rock crevices, flowers pale yellow, 
rare, K.M.Uou 9230, 17 Aug 1938 (holotype: PE). 

Specimens ex imined CHINA Gansu Kangxun on ua^ horn Y.ngba to Xiaoheba, in 
blitk anthers litquciit 9 Au^ 196^ / YZ/'nK /6/S^(PI ) 
14. Lycons traubii Ha> ward, PI Life M 40 1 95" Ii lub PI Life 13:44. 1957, and 

Lca\cs loiatc-lanccolate, toca 22 5 cm long, 1 2-2 1 cm broad, obtuse at 
apex, glabrous, not glaucous, with a distmct whitish stripe in the center. Scape 
to 50 cm long Spathe-vaKcs ovate, ^-5 cm long Pedicels 8-9 mm long. Fiow- 
(ismoico. Icsshoii/ontalh spuachng Pciigone i ic h orant^c-vcllow, with a 

Hsu, Synopsis of Lycoris 319 

deeper band in the center. Tepaltube 1.5-2 cm long, recurved downwards; cepals 
narrowly oblanceolate, 6.8-7 cm long, 1.3-1.6 cm broad, strongly recurved. 
Stamens slightly exserted. Style longer than stamens, reddish only on tip. 

Phenology: leaves appearing in autumn, about a month later than in L. aurea; 
scape produced in early September to October. 

Karyotype: 2n=lQU+2i: =12 {V>os>e 1958;Bose&Flory 1963; Kurita 1987b); 
2^;=9M+4T= 13 (Bose 1958; Bose & Flory 1963; Kurita 1987b); 2«=8]M+6T= 14 
(Kurita 1987b). 

Distribution: Taiwan and southernmost Japan, including southern Kiushu 
and Loochoo Islands. On slopes where moisture is sufficient and at edges of 
forests; to 100m. 

Specimens examined: JAPAN. Kagoshima Pref.: Ibusuki, S. Kurtta 891015 (CBM). Oki- 
nawa Pref.: Yonagunijima Isl., west of Sonai, K.Schimahukuro & Y.Miyagi 5198 (TI); Okinawa 
Isl., Kuganiimu, S.Kunta 931020 (CBM). 

CHINA. Taiwan: Keelung, S.lnariyarnas.n. (TI). 

Kuri ta( 1987b) detected two cytoraces having 2;^= 12 and 14 respectively 
with distinguishable morphological characteristics. The leaves of the race with 
2«= 12 are somewhat dark blue-green and the tepals are strongly recurved; in 
the race having 2n=l4 the leaves are lustrous yellow-green and the tepals are 
moderately recurved. Based on C-banding pattern, Kurita (1987b) suggested 
that the sterile cytorace having 2^= 1 3 may be a hybrid between the two fertile 
cytoraces having even chromosome numbers. 

15. Lycoris XalbifloraKoidz. (prosp.), Bot. Mag. Tokyo 38:100. 1924. Makmo, 
Acta Phytotax. Geobot. 13:18, pl.2. 1943; Traub & Moldenke, Amaryllidac: Tribe 
Amaryll. 178. 1949; Koyama, Baileya 7:4. 1959;Traub, PI. Life 22:59. 1966;Ohwi,H. 
Jap. ed. 2, 384. 1978; Hsu et al., Fl. Reipub. Pop. Sin. 16(1):22. 1985. Type: Not 
indicated. Koidzumi remarked that this species was cultivated in Japan and was perhaps 
spontaneous in the Amamiohshima Island of that country. 
Leaves up to 35 cm long, 1.2-1.5 cm broad, somewhat yellowish green with 
a rather inconspicuous whitish stripe in the center. Flowers pink in bud, open- 
ing creamy white, changing to white with age, with a very light orange-yellow 
stripe in the center of each tepal. Tepaltube ca. 2 cm long; tepals 6.5-7.5 cm 
long, 1-2.2 cm broad, moderately ruffled-margined, strongly recurved. Sta- 
mens long, far exceeding tepals. Style slightly exserted. 

Phenology: leaves emerging in autumn; scape produced in mid September to 
early October. 

Karyotype: 2;?= 5M+ 1T+ 1 1 A= 17 (Bose I960; Kurita 1987a); 
2n=5M+ 1T+ 1 1 A+ lm = 18 (Kurita 1987a). 

Distribution: southwestern Japan, mainly in Kyushu. In moist places by streams 
and hillsides, and disturbed places near human habitation and in graveyards; 
30-500 m. Also cultivated as an ornamental in Japan. 

320 SiDA 16(2) 1994 

Specimens examined: JAPAN. Kagoshima Pref.: Kaseda, Tojmbaru, H.Ohba &S.Akiyama 
2627 (TI). Kumamoto Pref.: Amakusa, Kawaura, M. Yamada 2643 (CBM). Okinawa Pref.: 
lejima Island, M.Tashtros.n. (TI). Shizuoka Pref.: Odawara, cult., S.Kunta s.n. (CBM). 

This Species is characterized by the creamy white flowers with long exserted 
stamens. It is somewhat comparable to L. radiata, but that species possesses 
crimson flowers with narrower tepals and narrower leaves, and leaves with a 
distinct whitish band in the center. 

The karyotype of L. Xalbiflora is very variable. Besides 2n= 17 & 18, comple- 
ments possessmg 2n= 16 (Inariyama 1931) and 19 (3M + 5T+ 1 lA) have been 
found in Japan and China. The cytotype having 2;^= 19 has been attributed to 
"L. aff albiflora Koidz." by Lin & Hsu (1989) and to "L. Xelegans" by Lin et al. 
(1990). L. Xelegans Liu & Hsu, ined. resembles L. aff. strammea Lindl. (Lin et al. 
1990, Xu et al. 1986,) or "L. sframmea" (Lm & Xu 1990) very much in gross 
morphology as well as in karyotype. This has caused much difficulty in identi- 

The origin of this highly sterile species with 2;?= 17 or 18 is problematical. 
Inariyama (1932, 1933, 1937) once considered it to be a hybrid between L. 
traubii (mistaken to be L. aurea) and L. sanguinea in the light of cytological point 
of view. Based on gross morphology, Makino (1943) supposed it to be a hybrid 
between L. radiata and L. aurea. Inariyama (1944), however, had changed his 
opinion and advocated strongly that the species originated from hybridization 
between L. traubii and L. radiata NUt.pumila. This was supported by Takemura 
(1962a), and especially by CaldweU (1981), who made crossing tests between 
the said species and as a result brought forth hybrids that were very similar to or 
almost exactly like L. Xalbiflora in gross morphology. But since L. radiata var. 
pumila does not occur in Japan, Kurita (1987a) suggested that the putative 
parents of L. Xalbiflora (5M+ 1T+ 1 1 A) were L. radiata (1 1 A) and L. traubii 
(5M+ IT). He argued that despite being a triploid sterile plant, L. radiata pro- 
duces some viable pollen occasionally (Koyama 1959). Moreover, the flowering 
period of the two species overlaps at times. The m type chromosome in the 
complement ofln^l^ might come from a race of L. radiata possessing such a 
chromosome together with the other 1 1 A type chromosomes. 

The origination of the cytotype 2?z=3M+5T+ 1 1 A= 19 ofL. Xalbiflora is in- 
teresting. This cytotype in China probably occurs only in cultivation. It has 
broader and less wrinkled tepals and shorter stamens. According to Kurita's 
(1987a) suggestion, it is ahybrid between two diploid species, one with 2n=l6, 
which produces gametes having 3M + 5T, and the other with 2«=22A, which 
gives rise to genomes consisting of 1 1 A. Based upon crossing tests, Lin et al. 
(1990) proved that both L. aff albiflora (2n=19) and L. aff. straminea are segre- 
gates in the Fi progeny of the L. haywardii X L. chinensis combination. 

Hsu, Synopsis of Lycoris 321 

(cult.), S.Caldwdl 549 (holotype: TRA); same locality, S.Caldwell 550, 55 1 (paratype: 
in Shanghai, China, in 1948. 

Leaves strap-shaped, 30—42 cm long, 1-1 .3 cm broad, rounded at the apex, 
deep green with a slightly distinct whitish midrib. Perigone creamy white, 
turning whitish with age, sometimes with reddish lines running along tepals or 
may develop a rose flush, with a greenish midrib underside. Tepaltube 8.5-12 
mm long; tepals linear-oblanceolate, up to 5 cm long, 8-10 mm broad, slightly 
ruffled-margined, recurved. Stamens exceeding tepals ca. 1/3, filaments creamy 
white, sometimes tinged with pink. Style exceeding the stamens, pink tipped. 

Phenology: leaves appearing in autumn, persisting to spring; scape produced 
in late July to August. 


Distribution: endemic to China (Zhejiang, known only in cultivation). 

Specimens examined: CHINA. Zhejiang: Hangzhou, Hangzhou Bot. Card., cult., Z.Z. Yu 
008 (HZBG). 

A specimen in PE collected in a garden called "Caojiahuayuan" in Shanghai 
without notes of collector and field number also belongs here. 

Kurita (1987a) suggested that this sterile species is a triploid hybrid between 
L. longituba, which produced the gamete having 3M+ 5T, and another species 
with an unreduced 22A, which was one of the following: L. radiata vzx.pumila, 
L. rosea, L. xhaywardii, or L. sprengeri. 

17. Lycoris stramineaLindl, J. Hort. Soc. London. 3:76. 1848, nomen subnud., 
and emend. Traub, PI. Life 12:42. 1956; Walp., Ann. Bot. Syst. 1:834. 1848-49; 
Kunth, Enum. PI. 5:546. 1850; Baker, Handb. Amaryll. 40. 1888; Spreng., Bull. Soc. 
Tosc. Ortic. 8:326. 1888; Worsley, Card. Chron. ser. 3,84:169. 1928; Traub & Mold- 
enke,Amaryllidac.: Tribe Amaryll. 178. 1949; Traub, PI. Life 13:43. 1957, in clavis; 
Hsu et al., Fl. Reipub. Pop. Sin. 16(1):18, pl.4, fig. 4. 1985. Type: CHINA. Robert 
Fortune 148 (holotype: MO). According to Traub (1956), this species was sent to the 
Kew Botanical Gardens by Robert Fortune from China in 1 845. The first author of the 
present paper, however, has been able to examine the type specimen of this species at the 
herbarium of Missouri Botanical Garden. The sheet (MO herb. no. 3149486) actually 

was based on the left hand one, while the right hand one was much slenderer in shape 
with narrower tepals (ca. 4 mm wide) and longer tepaltube (5.3-5.5 mm). 
Leaves strap-shaped, 24-49 cm long, 1.3-2 cm broad, obtuse at the apex, 
green with an inconspicuous whitish band in the center. Scape up to 22. 1 cm 
long. Perigone with tepals pale straw-colored, with a pink band and a few scat- 
tered red dots on the upper surface, changing to white in full blossom. Tepaltube 
4-5.5 mm long; tepals linear-oblong, 3.5-4.1 cm long, 5.2-12 mm broad, 
undulate margined and strongly recurved. Stamens exceeding tepals ca. 1/3. 
Style long exserted. 

Phenology: leaves appearing in autumn; scape produced in August. 

Karyotype: no reports. 

Distribution: endemic to China (Jiangsu and Zhejiang). In shady places in 
sparse woods; ca. 100 m. 

Specimens examined: CHINA. Jiangsu: Nanjing, Z.P. Wus.n. (JSBI). Zhejiang: Hangzhou, 
Feilaifeng, H.Q.Zhu 859 &1328 (HZBG); same locaiky, Hangzhou Bot. Gard., cuk. J.Z.Un 
001 (HZBG). No locality: Z. B. Wang 1 1 602 (PE). 

This species seems somewhat like L. Xalbifiora, but differs from that species 
in that the foliage appears in autumn, as well as by its smaller, pale-straw- 
colored flowers with pink stripes and red dots, and by its shorter tepaltube. 

This species may also be of hybrid origin, and the type specimens were prob- 
ably from segregates of the F] progeny of L. radiata var. pumila and a species 
with 2«= 16 combination. Lin et al. (1990) brought forth a hybrid between L. 
y^haywardii 9 and L. chinensis S , and they have named the hybrid L. aff 
straminea. It matches typical L. straminea in many respects in gross morphology 
and possesses 2«=3M+ 1 lT+5 A= 19- Their investigation has shown that as much 
as 81 .9% of the pollen of this sterile hybrid is shriveled. Besides, this hybrid not 
only failed to set seed under natural conditions, but failed to do so even under 
conditions of artificial pollination. 

18. Lycoris elsiae Traub, PL Life 14:43. 1958. Type: UNITED STATES. Tenne,s.see: 
Nashville, c\xk.,S.Caldwell 393 (holotype: TRA); same locality, S.Caldtvell594 (paratype: 

There are two specimens (W.H.PrestonJr. 964, 965) in MO (Herb. nos. 628, 629) trans- 
ferred from Traub Herbarium of rhe American Planr Life Society under the name of L. 

Leaves linear, 32-36.5 cm long, 1 .2-1.3 cm broad, rounded at the apex, dark 
green. Scape 30-70 cm long. Perigone soft salmon-colored, finally fading to a 
flesh color, with a deep pinkish band tinged with creamy and yellow along the 
center of each tepal. Tepaltube 1.2-1.3 cm long; tepals oblanceolate, ca. 4 cm 
long, up to 7 mm broad. Stamens exceeding the perigone. Style longer than 

Phenology: leaves appearing in autumn; scape produced in August to early 

Karyotype: 2n^\l (Bose I960; Bose & Flory 1963). 

Distribution: endemic to Japan. 

The identity of this Japanese species is not yet clear. It has scarcely been 
included in any published Japanese floras. It is morphologically somewhat com- 
parable to L. xhoudyshelii, but differs from that species in flower color and a 
longer tepaltube. It may be a hybrid between L. traubit and L. sanguinea var. 
kumana, both of which occur in the southern part of Kyushu. 

Hsu, Synopsis of Lycoris 323 

19. Lycoris radiata (L'Her.) Herb., Bot. Mag. 47:t.2113. 1821. Bot. Reg. 4, 
Append. 20, pi. 596. 1821; Hance,J. Bot. 12:262. 1874; Franch. &Sav., Enum. Pi. Jap. 

Bull. Soc. Tosc. Ortic. 8:326. 1888; Brecschneider, Hist. Eur. Bot. Disc. China 1:509. 
1898; Yashiroda, Card. Chron. ser. 3, 38:9, fig. 4. 1930; Traub & Moldenke, Amaryilidac: 
Tribe Amaryll. 177. 1949; Koyama, Baileya 7:2. 1959; Icon. Cormophyt. Sin. 5:549, 
fig. 792. 1976; Ohwi, FI. Jap. ed.2, 384. 1978; M.Kim & S.Lee, Korean. J. PI. Taxon. 

Nerine japonkaMvq., Ann. Mus. Bot. Lagduno-Batavum. 2:139. 1865-66. 

Lycoris terraccianii Dammann, CziAA'A. 1 889. 

Lycoris radiata (L'Her.) Herb. var. terraccianii Dammann, 1 .c. 
19a. Lycoris radiata var. radiata Type: not indicated. Baker (1988) remarked that this 
dens in 1750. The type material was most probably from China. 

Leaves narrow strap-shaped, up to 50 cm long, 3-8 mm broad, obtuse at 
apex, deep green with a whitish stripe in the center. Scape 30—60 cm long. 
Perigone bright red. Tepaltube 5-8 mm long; tepals narrow oblanceolate, 4- 
4.5 cm long, 5-6 mm broad, strongly crisped-margined and recurved. Stamens 
2-2.5 times the length of tepals. Style long exserted. 

Phenology: leaves appear in autumn and wither in April; scape produced in 
late September to early October. 

Karyotype: 2n=idA=^i{Bose 1959; Bose&Flory 1963; Fukudaetal. 1980; 
Inariyama 1931, 1933, 1937, 1951; Koyama 1962;Kurita 1987c; Lm& Hsu 
1989; Nishikawaet al. 1979; Nishiyama 1928); 2n= 1M+31A+ lm = 33 (Bose 
1963; Kurita 1987c); 2«=31A+ 1M' = 32 (Kurita 1987c). 

Distribution: 'jdipdiW (Aomori and southwestward), Korea, China, and Nepal. 
In moist, often disturbed places such as edges of paddy fields, margins of plan- 
Extensively naturalized in southeastern United States. 

Specimens examined: CHINA. Anhui: Chuxian, Langya Shan, Z.Z.Ding &J.S. Yue 0681 
iPEy,]inzhzuS.X.Sben 1101 (ACE); Huang Shan, L.G. Fa ^20 (JSBl); R.C.Chings.n. (NJU); 
Shucheng, East China Bot. Station 4620 (JSBI); Jixi, X.L.Liu 0^7 (PE). Fujian: Shanghang, 
L.G.Lin 6960 (VE);Anioy Univ. Fujian Exped. 1195 (PE); Nanjing, Amoy Umv. Fujian Exped. 
64564 (FUS); Ninghua, Fi/dan Univ. Fujian Exped. 91974 (FUS); Shaxian, Fudan Univ. Fujian 
Exped. 33473 (FUS); no locality, Y.Ltng 2583 (PE); Shucheng, East China Bot. Stat. 4620 (PE). 
Guangdong: Liannan, P.X. Tan 59386 (PE); Xinfeng, L.Deng8l47 (PE). Guangxi: no locality, 
Inst. Bot. Guangxi Exped. 3965 (PE). Guizhou: Jiangkou, Xuefeng Shan, on the south side of 
Mt. Fanjing Shan, Sino-American Guizhou Bot. Exped. 844 (PE); same locality, Daiyenfeng along 
the Kaitu River on the southwest side of Mt. Fanjing Shan, Sino-American Guizhou Bot. Exped. 
1133 (PE); Xishui, Bijie Exped. 1577,1712 (PE); Hexian, Anshun Exped. 1299 (PE). Hubei: 
Lichuan, R. Y.Dai &Z.H.Qian 920 (PE); G.X.Fu& Z.S.Zhang 1 745 (PE); Qianshih, C.C.Hos.n. 
(PE); Xuanan, H.J.Li 4662 (PE); Fangxian, K.M.Liou 8972, 9231 (PE); C.L.Cheng 509 (FUS); 
G.B.Hu 337 (FUS); no locality, PY.Li 5052 (PE); Zhuxi, P. Y.Li 9504 (SZ). Hunan: Baojing, 

324 SiDA 16(2) 1994 

LH.Uu9762 {PEy,Yizhiing,S.Q.Chen 1987 (PE,SZ).ymngsu:Nanimg,K.L.Chu 318iPBy, 
W.Z.Fang et al. 3^2 (SZ); Z.R. Wang 80 (FUS); X.C.Sun 5 (PE); Jiangning J.^, Yue 0561 (PE); 
Yixing, W.Z.Fang 196 (PE); Shanghai, Jiangwan, T.N. Yan 10412 (FUS). Jiangxi: Shangrao, 
S.S.Lat & M.X.Nie4750 (FUS, PE); Lushan, H.C.Cheo 98 (NJU); F.T.Wang s.n. (PE); K.C.Kuan 
74230 (PE); H.H.Hu2212 (PE); Nanchang, S.H.Hsimg 666 (PE); Anju, Wugong Shan,/ 5. 7//^ 
3283 (PE); Yingtan,7/^K^xi Mea'. £x^e^. s. n. (PE); Shangyou,y/^K^;c/ Ex/^e^. 0638 (PE); Jinggang 
Shan, 5.5.L^z 3023 (FUS); Xinning, KB.L/zo 3233 (PE). Shaanxi: way from Longwan to Shiquan, 
Sichuan: Xiushan, Xiushan Exped. 1136 (CDBI); Yaan, Yaan Exped. 1047 (CDBI); Rongjing, 
Rongjing Exped. 78-0724 (CDBI); Tianquan, Tianquan Exped. 78-134 (CDBI); W.P.Fang34l3 
(PE); K.C.Guan & W.T Wang 343 1 (PE); Hanyuan, Hanyuan Exped 0923 (CDBI); Eazu, Dazu 
Exped. 0838 (CDBI); Dazhu, Dazhu Exped. OSIS (CDBI); Xuanhan, Xuanhan Exped. 0467. 
0843 (CDBI); Kaijiang, Katjiang Exped. 0824 (CDBI); Daxian, Daxian Exped 0874 (CDBI); 
Youyang, Youyang Exped 1 133 (CDBI); Wulong, WulongExped. 1347 (CDBI); Pengshui, Pengshui 
Exped. 1104{CDBl)\M.t.'\,T.T-Yu23023 {V^)\G.H.Yang37300{VE)\C.W.Yao3121 {VE, 
SZ); W.P.Fang3381 (PE); 14867 (SZ);G.X.Xmg&K.Y.Lang 1712 (PE); W.P.Fangetal. 33431 
(PE); Y.H.Tao 31912 (SZ); Wuxi, G.H. Kzk^ 59502 (FUS, PE); Baoxing, K.L.Chu 3821 (PE); 
C.Pei 8243 (PE); Nanchuan, Jinfu Shan J.H.Xiong & Z.L.Zhou 93632 (PE, SZ); Guanxian, 
Qingchen Shan, Z.R. Wu 33936 (PE); Zhaohua, Y.Q.He 1 704 (PE); Jincheng Shan, Stchuan 
Econ. PL Exped. south group 89 (PE); Hefeng, H.J.Li 8018 (FE); Leshan, Z.TGuan 6301 (PE); 
Chongqing, Beibei, Z.L. CAo/^ <& Z.We i392 (SZ); Fengjie, Z.X.C;^o<&H,F.Z/^o» 26975 (SZ); 
Xuanen, H.J.Li 4662 (SZ); no locality, T.TYu 2363 (FUS); Chengkou, TR.Dai 102283 (FUS). 
Zhejiang: Lishui, S. Y.Chang 6336 (FUS, HZBG, PE); Panshan, T.N.Liou 8019 (PE); Siming 
Shan, Y YHo 27328 (HZBG, PE); 27746 (PE); Changhua, Y YHo 263 18 (HZBG, PE); Tianmu 
Shan, H.Q.Zhu 00377 (HZBG); Y.Y.Ho 23368, 263 18 (HZBG, PE); 23609 (HZBG); Zhepang 
PI. Resources Exped. 28860 (HZBG,PEy,YandangShan,S.G.Chen 461 (FUS); Pingyang, YYHo 
24736 (HZBG); D.X.Zuo etal. 24644 gSBI); Tiantai Shan, Y. YHo 27746 (HZBG); Suichang, 
Zhejiang PL Resources Exped. 26878 (HZBG); Longquan, Y.Y.Ho 21682 (HZBG); Jingning, 
Y.Y.Ho 24289 (HZBG); Hangzhou, S. Y.Chang 0639, 0837, 1202, 1448 (HZBG); Hangzhou 
Bot. Card., cult., J. Z.Lin 003, Z.Z. Yu 030 (HZBG). 

JAPAN. Chiba Pref.: Awaamatsu, Kiyosumi-yama, S.Kurita 84923 (CBM). Kanagawa 
Pref.: Hakone, T.Sawada 2340 (TI). Kyoto Pref.: Yagi-cho, Hunai-gun, G.Murata 19637 
(PE). MiyagiPref.:Sendai,Tomioka,H,H^raj.K. (TI). SagamiPref.: Enoshima,.^.y\lw//«e35^ 
(PE). Wakayama Pref.: Nishimuroo-gun, Tanabe-cho,/. Nakajima 13923 (TI); Ooita Pref., 
Yabakei, M.Togashi 743 1 (TI); Shiga Pref, Kohga-gun, Shigarakimachi, Asamiya, H.Ohba et 
aL 9048 (TI). Yamaguchi Pref.: Kuniyoshiki-gun, Oouchimura, Yada, S.Nikaido 107 (TI). 

This distinctive species is characterized by its very narrow leaves, bright red 
flowers possessing narrow, strongly crisped-margined and recurved tepals, and 
the very much exserted stamens and styles. 

This is probably the most widespread species ofLycoris. It occupies a large 
distribution area in China, mainly in the southern part of the Yangtze River. In 
Japan, it occurs everywhere except Hokkaido. The broad distribution of this 
sterile triploid is largely due to its strong vegetative reproduction-propagation 
carried on by the rapid formation of new lateral bulbs. 

But this triploid L. radiata var. radiata resembles the diploid ydx.pimiila Grey 
(see below) very much in gross morphology; they are difficult to differentiate 
one from another in the field or in herbarium. Accordingly, many of specimens 

Hsu, Synopsis of Lycoris 325 

under the name L. radiata may actually be L. radiata 'VAi.pumila. The ecological 
preference of these two taxa, however, is quite different: L. radiata var. radiata 
grows usually in disturbed habitats such as edges of paddy fields, margins of 
plantations, waste places around dwellings, and graveyards, whereas L. radiata 
V2x.pumila grows in more natural habitats such as open slopes, shaded places by 
streams, sparse woods, etc. 

With regard to the origin of this triploid L. radiata (2;z=33), the most com- 
mon way would be the hybridization of a diploid with a tetraploid. But because 
tetraploids have never been found in this species, Liu & Hsu (1989) suggested 
that L. radiata originated from the combination of an unreduced gamete of a 
diploid with a normal gamete of another diploid of L. radiata \2s.pumila {2n=22). 

This sterile species was supposed to be an autotriploid (Inariyama 1951; 
Nishiyama 1 928). Kihara & Koyama (1 954) indeed obtained triploid offspring 
of L. radiata by self-pollination. But Kurita (1987c) argued that it is structur- 
ally heterozygous at least with regard to the satellite chromosomes and the four 
chromosomes carrying rather smaller short arms so far as somatic karyotype is 

Koyama (1959), Kurita (1987c), and Maekawa (1943), suggested strongly 
that L. radiata var. radiata must have been originated in China and was intro- 
duced into Japan by people. The diploid L. radiata NZi.pumila occurs only in 
China and has never been found in Japan and elsewhere. L. radiata var. radiata 
has long been known as a hardy plant since ancient time, and usually occurs in 
habitats disturbed by people in both China and Japan. In America, it is culti- 
vated outdoors in the southeastern states and on the Pacific coast. 

A cultivar with white flowers has been reported. There is a picture of this 
cultivar in Creech's (1952) paper. Recently, a new form of L. radiata var. radiata 
was proposed by Yonezawa (1989): forma bicolor, the tepals of which were rose- 
colored and white-margined. The type specimens of this new form were all 
collected from Kyoto City of Honshu, Japan. 

19b. Lycoris radiata (LHer.) Herb. ^ 

Traub & Moldenke, Amaryllidac: Ti 

This fertile diploid taxon resembles var. radiata in external morphology and 
is very difficult to discriminate from it. So it is quite evident that many speci- 
mens cited under L. radiata var. radiata actually belong here. 

Karyotype: 2«= 22A= 22 (Bose 1 958; Hsu et al. 1 984; Inariyama 1951; Koyama 
1962; Kurita 1987a; Liu & Hsu 1989; Nishikawa et al. 1979; Takemura 1962a; 
Yoshida 1972). 

Distribution: endemic to China (Shaanxi, Henan, Anhui, Jiangsu, Zhejiang, 
Jiangxi, Fujian, Guangdong, Guangxi, Hunan, Hubei, Sichuan, Guizhou, and 
Yunnan). Often in open moist slopes, shaded wet places by streams, sparse woods, 

and sandy flood lands; 50-1000 m in eastern, central, and southern China; to 
2500 m in southwestern China. 

This variety hybridizes with many other species ofLycoris and gives rise to 
fertile hybrids. According to Kurita (1987a), hybrids between this variety and 
L. sprengeri have been called "L. cv. Sprenpumila" and are well known to be fully 
fertile in spite of the morphological disparity of the parental taxa. Hybrids be- 
tween L. radiata yax.pumila and L. chinensis, L. longituba, or L. aff. strammea, have 
various rates of fertility (Lin et al. unpub.). 

19c. Lycoris radiata (L' Her.) Herb. var. kazukoana Yonezawa, J. Phytogeogr. 

Tax. 37(2):73, fig. 1, 1989. Type: japan. Kyoto Pref.: Honshu, Matsugasaki, 

Sakyoku, Kyoto City, K. Yonezawa & N. Yonezawa s.n. (holotype: in Kana no. 1 30006); 

same locality, N.Yonezawa 10100, 10120 (paratypes: in Kana nos. 130005, 130006). 

This variety differs from var. radiata in its flowers being smaller and pale red 

or becoming whitish in color, and the tepals 1.5-3.2 cm long and 2-5 mm 

broad, only slightly reflexed. 

Phenology: scape produced in late September to October. 

Karyotype: no reports. 

Distribution: endemic to Japan; 70-250 m. 

20. Lycoris XroseaTraub & Moldenke (pro sp.), Amaryllidac: Tribe Amaryll. 
178. 1949. Hsu et al., Fl. Reipub. Pop. Sin. 16(1):20, pl.4, fig. 5-6. 1985. Type: 
CHINA' Not cited or mention by Traub and Moldenke (I.e.), 

Leaves strap-shaped, up to 44 cm long, 7-1 1 mm broad, rounded at apex, 
light green with an indistinct whitish stripe in the center. Scape 30-60 cm 
long. Perigone rose-colored. Tepaltube 1-1.2 cm long; tepals oblanceolate, 4-6 
cm long, 7-8 mm broad, ruffled at the base, slightly recurved. Stamens exceed- 
ing tepals ca. 1/6. 

Phenology: leaves emerging in autumn; scape produced in September. 

Karyotype: 2«= 22A = 22 (Hsu & Huang 1984; Liu & Hsu 1989). 

Distribution: endemic to China (Jiangsu). In sparse second growth forest of 

Specimens exammed: CHINA. Jiangsu: Shanghai, Sheshan, G.J. Fan s.n. (PE); same locality, 
Hengshan, Y. Hsu et al. 63918 (SHMI). Zhejiang: Hangzhou Bot. Gard., cuh. J.Z.Un 

This species is characterized by its rose-colored flowers and by its stamens 
only 1/6 longer than the tepals. It is somewhat comparable to L. radiata var. 
radiata and wzt-pumila, but in those taxa the flowers are bright red and the tepals 
are narrower and strongly crisped-margined and recurved. 

Lycoris Xrosea is most probably a natural hybrid. From the karyoiogical point 
of view, Kurita is of the opinion that this species must be the hybrid of L. 

Hsu, Synopsis of Lycoris 

radiata v^t.pumila (2^=22A) and L. sprengen {2n=22A). His ad^ 
ported by Lin et al. (unpub.), who have made a crossing betwet 
and as a result have brought forth a hybrid that is fertile and ; 
gross morphology with L. Xrosea. 

Lycoris albiflora Koidz. cv. Lactiflora 

Belongs to L. Xalbiflora in its broad sense. 
Lycoris "cinnabarina" (sphal. "cinnabarinum" a "catalog" name) 

This species, reported only in cultivation, is very similar to L. sangmnea Maxim. 
Hunt (1963) referred it to be L. sanguinea var. cyrtanthifiora Hort. Caldwell (cf. 
Easterly 1969) said that it might turn out to be L. kiusiana Makino (= L. san- 
guinea Maxim, var. kiusiana (Makino) Koyama). Easterly (1969) reported the 
karyotype of L. "cinnabarina" to be 2«=21A+ IM, slight different from that of 
L. sanguinea. Williams (1983), however, suggested that it was most likely to be 
a hybrid between L. sanguinea X L. traubii, with a karyotype of 2?2= 14T+4M= 18. 
Lycoris Xelegans Lin, Yu, & Hsu in S.A.He et al. (ed.), Proc. Intern. Symp. Bot. 
Gard. 565. 1990, nom. nud. =L. sprengen 9 X L. chinensis 6 belongs to L. 
Xalbiflora in its broad sense. 
Lycoris flavescensU.Kim & S.Lee, KoreanJ. PI. Taxon. 21:127. 1991- 

This species is probably a hybrid between L. sanguinea var. koreana and L. 

;. 47:t.21 13. 1819 = Griffinia hyactnthina 

Lycoris Xjacksoniana Traub, PL Life 20:52. 1964 = L. sprengeri x L. radiata (prob- 
ably var. pumila) 
Lycoris josephinae Tmuh, PL Life 21:63- 1965, and 22:60. 1966. 

The type material {Traub 792, holotype) was collected in Sichuan, China. 
According to the type description, there seems little difference in morphologi- 
cal characters deserving specific consideration between L.josephinae and L. ra- 

Lycoris xlajollaTtz.\xh, PL Life 19:50. 1963 = L. aurea x L. traubii. 
Lycoris radiata Miq., Ann. Mus. Bot. Lugduno-Batavum 2:139. 1865-66 = 
Ungernia trisphaera Bunge. 

Lycoris sewerzowii Kegel, Bull. Soc. Not. Moscou. 41:435. 1868 = Ungerma 
sewerzawii (Kegel) Fedtsch. ex Vvedensky. 
Lycoris Xwoodii Traub & Moldenke, PL Life 13:85. 1957 = L. radiata 9 x 

We thank the curators of the following herbaria for providing loans or allow- 
ing access to collections: Institute of Botany and Kunming Institute of Botany 
of Chinese Academy of Sciences, Sichuan University, Anhui Normal University, 
Jiangsu Institute of Botany, University of Tokyo, and Missouri Botanical Gar- 
den. We are especially grateful to Dr. Mikio Ono and Mr. Y.M.Yu of Makino 
Herbarium of Tokyo Metropolitan University for sending us difficult-to-obtain 

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loid species. Cytologia 52:137-149. 

Intrapopulational and/or intraspecific variation in the karyotype of L. sanguinea Maxim, var. 
khniana and L. sanguinea Maxim, var. koreana (Nakai) Koyama. Cytologia 53:307-321. 
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of karyotype evolution in the genus. Cytologia 53:323-335. 

330 SiDA 16(2) 1994 

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Botanical Research Institute of Texas 

The genus Cyhianthus Martius contains ten subgenera and perhaps 150 spe- 
cies (Pipoly 1987, 1991, 1992). Subgenus C)/^i^)^^/?^j contains approximately 
52 species, including 7 Peruvian ones as yet undescribed. The subgenus is dis- 
tributed in the Andes from Colombia to Bolivia, thence eastward through the 
Guayana Floristic Province (sensu Maguire 1979) to French Guiana and south- 
eastward through Amazonia to southeastern Brazil. The subgenus is defined by 
a vestigial pistillode in staminate flowers, basifixed anthers as wide or wider 
than long, and a staminal tube which is developmentally fused to the corolla 
tube; the stamens thus appearing epipetalous. 

During phytodiversity studies in the Amazon Basin of Colombia, concomi- 
tant with floristic and monographic studies of the genus Cyhianthus for Flora 
Neotropica and Flora de Colombia, I have encountered the following new species, 
described herewith. 

Cyhianthus barbosae Pipoly, sp. nov. (Fig. 1) 

Propter ramulos angulatos rufo-stellato-tomentosos, folia pseudoverticillata, lobos calycinos 
late ovatos vel suborbiculares ad apicem rotundatos vel truncacos, antheras latiores quam longiores 
ad apicem obtusas vel truncatas C. venezuelanum arete affinis, sed ab eo ramulis dense rufo- 
stellato-tomentosis et gianduloso-lepidotis (non solum rufo-stellato-tomentosis), laminis 

mm longo, corolla chartacea (non carnosa) 1.8-2.5 (nee 1.6-1.8) mm longa, coroUae tubo 

Shrub or small tree to 2 m tall; growth following Rauh's architectural model. 
Branchlets angulate, 3^ mm diam., with prominent, narrow longitudinal ridges, 
densely rufous-stellate and translucently glandular-lepidote, the tomentum per- 
sistent. Leaves pseudoverticillate; leaf blades membranaceous, linear-lanceolate, 
very narrowly oblanceolate or rarely oblong, (lO-)l 5-22(-27) cm long, 2-3-5 
cm wide, apex long-attenuate or acuminate, the acumen 1.5-2.5 cm long, base 
long attenuate and cuneate, decurrent on the petiole, the blade bullate when 
fresh, subbullate when dry, nitid, minutely and sparsely orange-punctate above, 
below densely pellucid-punctate, sparsely black-punctate-lineate and with scat- 
tered stellate trichomes below, venation brochidodromous, costa slightly raised 
above, prominently raised below; secondary veins 18-24 pairs, deeply impressed 
above, prominently raised below, the margin irregular, undulating, flat; petioles 
marginate, 0.5-1 cm long, sparsely stellate above, densely stellate below, slightly 
pulvinate at base, glabrescent. Staminate inflorescence a simple, erect raceme 8— 
1 1 cm long at maturity, the rachis and pedicels densely rufous-glandular-granu- 
lose and rufous-stellate-puberulent; inflorescence bract unknown; floral bracts 
membranaceous, Imear, (1.5-)2-2.4 mm long, 0.2-0.3 mm wide, equal to or 
longer than the pedicel, apex subulate, glabrous above, densely stellate below, 
carinate, the margin entire, with scattered stellate hairs; pedicel cylindrical, 
thin, (1 .5-)2-2.3 mm long, densely glandular-granulose and with a few scat- 
tered stellate hairs, glabrescent. Flowers chartaceous, 4-merous, green, pendent 
at maturity; calyx 1.2-1 .4 mm long, deeply divided, tube 0.3-0.4 mm long, 
lobes very widely obovate to suborbicular, symmetric, 0.9-1 • 1 mm long, 0.8- 
1 . 1 mm wide, translucent, apex broadly rounded to truncate, thickened medi- 
ally, densely and prominently orange-punctate, abruptly constricted basally, the 
margin white to hyaline, irregularly erose, sparsely glandular-ciiiate; corolla 
subrotate, 1 .8-2.5 mm long, translucent, the tube quadrate in cross-section, 1- 
1 . 1 mm long, densely rufous-glandular-granulose and rugose without, espe- 
cially on the surface alternate with the calyx lobes, densely translucent-glandu- 
lar-granulose within on surface alternate with anthers, the lobes depressed-ovate, 
0.8-1 .4 mm long, 1.8-2.2 mm wide, apex very broadly rounded to truncate, 
with at least one apical notch, densely and prominently orange-punctate and 
rufous-glandular-granulose without, densely translucent glandular-granulose 
within except near base; anthers widely triangular, apparently sessile and epipe- 
talous (the staminal tube not easily discernible from corolla tube), 0.4-0.6 mm 
long, 0.8-1 mm wide, apex obtuse, base truncate, appearing truncate at anthe- 
sis, birimose, dehiscent by large terminal pores opening into wide longitudinal 
slits for ca. 3/4 anther length, connective adnate to corolla tube adaxially, densely 
red-punctate abaxially; pistillode conic, 0.3-0.6 mm long, 0.2-0.4 mm diam.. 

Type: COLOMBIA. Amazonas. Municipo de Leticia: Parque Nacional Natural Amacayacu, 
Quebrada Agua Pudre, ca. 1.5 km NE of Quebrada mouth at rio Amacayacu, permanent 25- 
hectareplot, 03°47'S, 70°15'W, 200-220 m, 18 Nov 1991 (stam. fl) J. Ptpoly et al. 16450 
(holotype: COL; isotypes: BRIT, MO, K, TEX, US). 

Paratypes: COLOMBIA. Amazonas. Municipio de Leticia: Parque Nacional Natural 
Amacayacu, Quebrada Agua Pudre, ca. 1.5 km NE of Quebrada mouth at rfo Amacayacu, 
permanent 25-hectare plot, 03°47'S,70°15'W, 200-220 m, 18 Nov 1991 (ster.),/ Pipolyetjl. 
15790 (COL, FMB, MO); (ster.),/ Pipolyetal. 16509 (BRIT, COL, FMB, MO, TEX); (stam. 
fl),7. Pipoly etal. 16513 (BRIT, COL, F, FMB, MO, TEX); (stam. fl),/ Pipo/y 16528 (BRIT, 

Distribution, ecology and conservation status: Cybianthus barbosae grows in tall wet 
forest on lateritic soils and is presumably endemic to the Leticia area. The canopy 
of the forest in which it occurs is approximately 35 m in height, and the domi- 
nants include Virola, Osteophloem and Iryanthera (Myristicaceae), Eschweilera 
(Lecythidaceae), Licania (Chrysobalanaceae), and Parkia (Mimosaceae). Cybianthus 
barbosae grows specifically along small streambanks on rotten logs, at 200-220 
m elevation. In the area of a quantitative inventory in the park, I observed six 
individuals of C barbosae per hectare, a relatively high frequency for a species of 

Etymology: This species is dedicated to Cesar Eduardo Barbosa Castillo, Direc- 
tor of the Herbarium, Unidad Investigativa Federico Menem (FMB), Mininstry 
of Environment, Colombia. Cesar Barbosa is a specialist in Pithecellobmm and 
Rhizophoraceae of Colombia, and a keen student of the Amazonian flora. He 
first spotted Cybianthus barbosae growing next to a 40 meter tree oiSterigmapetalum 
(Rhizophoraceae) sp. nov. 

Widely ovate or suborbicular calyx lobes, rounded or truncate apically, an- 
thers as wide as long and obtuse or truncate at apex, and rufous-stellate-tomen- 
tose, angulate branchlets indicate that Cybianthus barbosae is most closely related 
to C. venezuelanus , a taxon dispersed from Guyana west through Amazonian 
Brazil and Venezuela to Colombia, thence south through the Andes to Peru. 
The widely trangular anthers, quadrate corolla tube, glandular granules inter- 
spersed with rufous stellate hairs on the branches, marginate and somewhat 
pulvinate petioles, and the chartaceous and longer perianth, immediately sepa- 
rate C. barbose from C. venezuelanus. 

Cybianthus ruforamulus Pipoly, sp. nov. (Fig. 2) 

', Notes on the genus Cybia 

338 SiDA 16(2) 

Tree to 8 m tall; growth follow 
portions densely rufous hirtellou 
just above base (Y-shaped), stiff, persistent. Branchlets terete, 8-10 mm diam. 
Leaves alternate; leaf blades chartaceous, oblanceolate, (34-)38-46(-52) cm long, 
(9-)l 1-13(-16) cm wide, apex acute to short-acuminate, the acumen 0.3-0.5 
cm, base long-attenuate and cuneate, decurrent on the petiole, subbuUate, sor- 
did above, pallid below, densely prominently and minutely black punctate above 
and below, more densely tomencose along the costa and veins below, venation 
brochidodromous, the costa impressed and hirtellous-tomentose above, promi- 
nently raised and densely hirtellous-tomentose below, secondary veins (1 5-)20- 
24 pairs, deeply impressed above, prominently raised below, the margin entire, 
flat; petiole deeply canaliculate, pulvinate, 2-3 cm long, densely tomentose, 
persistent. Staminate inflorescence a simple, erect raceme 8-10 cm long, the 
rachis, pedicels, and calyx densely tomentose; inflorescence bract unknown; floral 
bracts linear, 1.3-1.5 mm long, 0.2-0.3 mm wide, apex subulate, hyaline, densely 
long-hirsute abaxially, glabrous adaxially, margin entire; pedicel cylindrical, 0.4- 
0.6 mm long, densely tomentose, persistent. Flowers chartaceous, 4-merous, 
greenish-maroon, translucent, pendent at maturity; calyx cotyliform, deeply 
divided, 1 . 1-1 .4 mm long, tube 0.2-0.3 mm long, lobes widely ovate to subor- 
bicular, 0.9-1 • 1 mm long, 0.7-0.9 mm wide, apex obtuse, densely and promi- 
nently red punctate, the punctations crowded and appearing vermcose throughout 
except at scarious margin, tomentellous alternating with the calyx lobes, mar- 
gin irregular, entire, bifid-hirtellous; corolla subrotate, 1.3-1.7 mm long, the 
tube 0.3-0.5 mm long, the lobes depressed-ovate 0.9-1.2 mm long, 1.3-1.5 
mm wide, apex broadly rounded, not notched, densely and prominently red- 
punctate and appearing verrucose and tomentose on areas between calyx lobes 
abaxially, densely glandular-granulose throughout within, the margin entire, 
hyaline, tomentose; anthers subsessile, subdeltate, 0.3-0.4 mm long, 0.4-0.5 
mm wide, apex acute, base truncate, apically dehiscent by pores, the pores not 
birimose, opening ca. 3/4 anther length, the connective densely and promi- 
nently punctate dorsally; pistillode lageniform to obturbinate, 0.5-0.7 mm long, 
0.1-0.2 mm, glabrous, style short, stigma subcapitate. Pistillate inflorescence: 
like the staminate but 6-9 cm long. Fruit sessile, calyx 1.3-1.5 mm long, the 
tube ca. 0.1 mm long, the lobes widely ovate, 1.2-1.3 mm long, 0.7-0.9 mm 
wide, apex obtuse, margin as in staminate. Fruit globose, 8-10 mm diam., 
orange, then purple, then black at maturity, rugose, inconspicuously black- 
punctate, exocarp thick, juicy. 

TYPE: COLOMBIA. Amazonas: along rio Yari, near mouth of Quebrada El Mochilero, ca. 
00°30'N, 72°53'W, 120-200 m, 23 Apr 1986 (sram. fl), G. GaleamJ. H. Torres,]. Huitoto, & 
B. Plazas 1103 (holotype: COL). Fig. 2. 

Paratypes: COLOMBIA. Amazonas: Quebrada Aduche, 200 m, 5 Aug 1977 (fr) J, M, 
Idrobo 8908 (COL). BRAZIL. Acre: Estrada Alemanha, Cruzeiro do Sul, 14 Apr 197 1 (fr), G. 

PiPOLY, Notes on the genus Cybianthus 339 

Pranceetal. 11908 (IAN, INPA, K, MG, NY); 6 May 1971 (fr), P. Maas etal. P12737 (¥, IAN, 
INPA, K, NY). 

Distribution, ecology and conservation status: Cybianthus ruforamulus is known 
from terra firme forests, near 200 m, in Amazonas, Colombia and Acre, Brazil. 
According to one collector (Galeano, pers. comm.), it grows along small brooks 
and other minor watercourses in primary forests, and may be considered an 
indicator of environmental quality. 

Etymology: The specific epithet describes the rufous tomentum of the branchlets, 
composed of forked trichomes unique within the genus. 

Cybianthus ruforamulus appears to be closely related to C. minuttflorus,2i poody 
known taxon from the rim of the Amazon Basin in Peru, but is easily recognized 
by the biramose-hirtellous tomentum of the branchlets, leaves and inflorescence, 
the shorter petioles, the chartaceous, greenish-maroon perianth, and the sub- 
deltate, glabrous anthers. The tomentum of Y-shaped hairs (Fig. 2C) is the first 
of its kind reported for the genus and should not be confused with the mal- 
pighiaceous trichomes unique in Cybianthus to subgenus Triadophora. 

Research for this paper was completed at the Missouri Botanical Garden, 
where my studies in Amazonian biological diversity were supported by grants 
from the John D. and Catherine T. MacArthur Foundation. Cybianthus barbosae 
was found during my course in forest inventory and tropical tree architecture 
offered to graduate students of the Universidad Nacional de Colombia, Insti- 
tuto de Ciencias Naturales, in Parque Nacional Natural Amacayacu. I thank 
my Colombian counterpart, Agustfn Rudas LI. for logistical support. I also owe 
special thanks to Oscar Pinto, former regional director of INDERENA, Cesar 
Barbosa, Jaime Aguirre, Clara Ines Orozco, Pilar Franco, Adriana Prieto, Pablo 
Palacios, Santiago Duque, Jose JVlurillo, Roscio Cortes, Marta Gonzalez, Roberto 
Sanchez, and Hector Esquivel, for attending the course or facilitating arrange- 
ments. Linda Ellis executed the fine line drawings of the taxa. 


Maguire,B. 1979.Guayana,reg 

on of the Roraima Sandstone Formatio 

. In; K. Larse 


L.B. Holm-Nielse 

n, eds. Tropi 

ai Botany Academic Press. London. Pp 



laceae. In: A 






ematic revis 

on of the ge 


rmdenia (Bentham) 


le). Mem. N 

ew York Bo 

t.Gard. 43:1-76. 




rpha (Myrsin 


100? Th 

e genus Cjfo 

.nthus. nhg 



Missouri Bot. Gar 

. 79:908-957. 







Botanical Research and Consulting 

P.O. Box 67 17 
Bryan, 7X77805-6717, U.S.A. 

jnty, Texas. A key to the species ot sect 
n Phaestoglochin, is considered distinct. 

Carexperdentata, seccion Phaestoglochin del con 
. las especies de la seccion Phaestoglochin que v 
. Se lectotipifica la seccion Phaestoglochin de 

Carex section Phaestoglochin [Sy = section Bracteosae sensu Mackenizie (193 1), 
non Pax}, subgenus Vignea (P. cie Beauvois ex Lestiboudois f.) W. Petermann was 
described from Europe by Dumortier in 1827. This section now includes 27 
species. However, most of the taxa are native to the New World. Today, all of the 
European species have been introduced into North America. Their distribution 
in the New World is primarily in temperate North America. However one 
species, C. xalapensis C. Kunth, is found in Mexico and northern Guatemala. 
Section Phaestoglochin is characterized by 1) an inflorescence with 3-25 sessile 
spicate branches, frequently with ten or less branches, the lower branches occa- 
sionally being compound in certain taxa (especially C. muehlenbergii C. Schkuhr 
ex C. Willdenow var. enervis W. Boott); 2) spikes androgynous, rarely pistillate, 
or having short staminate spikelets arising laterally from an androgynous or 
pistillate spike; 3) perigynia plano-convex or unequally biconvex, with the bod- 
ies of the perigynia more or less abruptly contracted into a beak; 4) achenes two- 
sided, either lenticular or slighdy biconvexed; 5) style jointed with the achene; 
6) stigmas two. Since section Phaestoglochin has never been typified, C. muricata 

L. is here designated as the type for section Pbaestoglochin. The typification of C. 
muricata has been reviewed by Reznicek and Ball (1980). 

Section Bracteosae R Pax differs from section Pbaestoglochin by 1) a denser 
inflorescence with more spikes; 2) perigynia with a spongy area at the basal end 
of the ventral surface, occasionally on the dorsal surface as well; 3) and usually 
with wart-like bumps over the lower part of the ventral surface and sometimes 
across the dorsal surface of the perigynia. Kunth (1837) was the first to use the 
name Bracteosae as an infrageneric category, but without rank or description. Pax 
(1889) published Bracteosae at the sectional rank and provided a scant and am- 
biguous, but valid description. Pax did not cite Kunth, giving authorship of 
section Bracteosae solely to himself Pax cited Carex cephalophora Muhlenberg ex 
Willdenow of North America as belongmg to this section. This is undoubtedly 
where Mackenzie (1931) got his concept that the North American taxa that he 
treated belonged to section Bracteosae. Klikenthal (1909) recognized the North 
American taxa that Mackenzie (1931) recognized as section Bracteosae as section 
Mulhlenbergianae, a superfluous name for section Pbaestoglochin, and recognized 
the South American taxa as belonging in a section distinct from those in North 

While researching the following complex of species in section Pbaestoglochin; 
C. austrina Q. Small) K. Mackenzie, C. cephalophora H. Muhlenberg ex C. 
Willdenow, C. leavenwortbit C. Dewey, C. mesochorea K. Mackenzie, C. mueblenbergii 
Schkuhr ex Willdenow var. enervis W, Boott, and C. mueblenbergii C. Schkuhr ex 
C. Willdenow var. mueblenbergii, a new species oi Carex (C. perdentata S.D. Jones) 
was discovered. 

Means, variances, standard deviations, and ranges were measured for nine 
populations consisting of 10 plants per population. Additional specimens were 
examined but no measurements were found to lie outside of the range recorded 
for the nine populations. The mean, plus and minus two standard deviations, 
for all parametric data is presented in the species description. If the ranges ex- 
tend beyond the mean, plus or minus two standard deviations, then the exten- 
sion is listed as parenthetical. Fruiting dates are based on a statistical mean, plus 
and minus two standard deviations. Ranges greater than and/or lesser than the 
two standard deviations are listed as parenthetical. 

Carex perdentata S.D.Jones, sp. nov. (Fig. 1) 

(18.0-)20.6-46.4(-55.5) cm longis, 2.2-4.5(-4.6) mm latis; paginis adaxialibus papillosis; 
vaginis arctis. Inflorescentiis (5-)6-12(-l4) androgymis spicis, 1 3.5-25.3(-28,0) mm longis. 

344 SiDA 16(2) 1994 

Hahit perennial, cespitose; rhizmm short to long, dark brown to black, fibrillose; 
fertile culms {11. Q-)26M-lA.2{-90.{)) cm tall, erect, stiff, 2.0-3.2 (-3.8) mm 
wide ca. 2 cm above rootstock, sharply triangular, scaberulous above, leafy on 
lower third, conspicuously exceeding leaves; leaves with well-developed blades 
(4-)5-9 per fertile culm; hypostomous; blacks (1 8.0-)20.^46.4(-5 5 .5) cm long, 
2.2^.5M.6) mm wide, erect-ascending to slightly arching, thick, light green, 
flat, long-acuminate, margins antrorsely serrulate; abaxial surface v^lthont papil- 
lae or papillose distally; midvein raised, becoming increasingly more antrorsely 
scabrous distally; epidermal cells rectangular and raised above the braided pat- 
terned cell walls; adaxial surface papillose, papillae arising perpendicular from 
distal end of embedded clavate shaped epidermal cells (Fig. 2); stoniata narrowly 
elliptic-oblong, paracytic with subsidiary cells mostly triangular with some semi- 
circular in shape; sunken below adjacent epidermal cells, restricted to intercos- 
tal zones on abaxial surface; sheaths tight around culm; dorsal sheath surface not 
septate-nodulose or rarely inconspicuously septate, papillose, pale green; ventral 
sheath surface whitish-hyaline, striate, papillose (at least proximally), deeply con- 
cave and more or less callused at apex of distal end, yellowish-brown tinged; 
ligule 0.4-1 .9(-2. 5) mm long, membranous, more or less linguiform, or infre- 
quently acute at apex; inflorescences of (5-)6-12(-l4) androgynous spikes, 13.5- 
25.3(-28.0) mm long, (9.0-)9.5-l4.5 mm wide, usually 0.5-2. 5(-3) times as 
long as wide, but occasionally as wide as long; lateral spike second from bottom 
(4.5-)4.7-7.9(-8.0) mm long; stamtnate flowers few, with ovate-lanceolate cus- 
pidate scales; lowest inflorescence bract 7 .0-56.6(-l 05 .0) mm long including awn, 
pistillate scale-like but with larger and conspicuously longer awns; upper bracts 
pistillate scale-hke but with longer awns, mid-point of awns 0. l-0.5(-0.9) mm) 
wide; lateral Irranch , second from bottom (spike) (4.5-)4.7-7.9(-8.0) mm long; 
gap (internode) between the lowest two spikes (0.5-)0. 7-3. 5(^.2) mm\ pistil- 
late scales {1 .6-)l .S-i . I mm long, 1.4-2.0(-2.2)mm wide, 1-veined, rarely 3- 
veined, ovate, brownish or greenish-hyaline, narrower than and (excluding awn) 
about length of bodies of perigynia, apex usually conspicuously awned (rarely 
acuminate), awn (0-)0. 1-3.6 mm long , green mid-stripe, 0.2-0.4(-0.5) mm 
v^ide; perigyma (3.3-)3.4-5.2(-5.6) mm long, 1.9-2.7(-2.8) mm wide, (2-)4- 
19(-24) per spike, ascending or at maturity spreading, flattened-plano-convex, 
ovate, round-tapering and often slightly spongy at base, ventral perigynia sur- 
face veinless, or l-5(-8) fine veins, dorsal perigynia surface veinless or 1-10(- 
1 1) fine veins, submembranous, sharp-edged to base, serrulate above middle, 
tapering or abruptly contracted into a serrulate beak, beak bidentate, sutures 
conspicuous; teeth ( 1 .0-)l .4-1 .7(-l .8) mm long, narrowly-triangular; achenes 
1.8-2.6(-2.8) mm long, 1.5-2.1(-2.2) mm wide, lenticular, strongly flattened, 
ovate (Fig. 3) to suborbicular, substipitate 0. 1-0.3 mm long, minutely apicu- 
late, occiisionally retuse at apical end; epidermal cells are nonisodymetric; single 

large central silica body and no satellites perched on edge of silica platform (Fig. 
4); central body occasionally composed of two fused bodies; style straight, short, 
slender, enlarged at base, jointed with achene; stigmas two, reddish-brown, short; 
anthers ( 1 . 3-) 1.4-1. 8(- 1 . 9) mm long; chromosome number vin\^nov^fn\ fruiting ( 1 3 
Mar-) 21 Mar - 1 Jun (-12 Jun); ecology a facultative sciophyte, primarily in 
alfisols or inceptisols with sandy or sandy loam soil, sandstone outcrops, granitic 
outcrops, or thin soil over limestone, open mesic to submesic hardwood forests, 
or open hardwood-juniper forests, or woodlands in savannas in granite outcrops; 
elevation 175-525 m; dtstnbutton (Fig. 5), central Texas norrh to Oklahoma; 
economic importance forage value for livestock is low in palatability but is of use for 
wildlife, especially for rabbits, rodents, deer, and birds; it is also important in 

Typus: U.S.A. Texas. Palo Pinto Co.: 2.3 mi S on FR 4 from its jet. with FR 3137, S of Palo 
Pinro, mesic to submesic live oak dominated W-facing slope with reddish sandy loam soil and 
sandstone, 23 Apr 1992, S. & G. Jones 8349 (holotype: MICH; isotypes: BRIT, MO, OKL, 

Additional specimens: OKLAHOMA. Comanche Co: 1 .3 mi N of Meets along W side of 
Rt. 115, lightly shaded, moist soil along trail through deciduous forest, frequent, 15 May 
1 990, Naczi 2414 B (MICH). Cleveland Co.: low, damp, ground, 28 May 1939, Bebh4085 

' (BRIT/SMU, ctb, MICH, MO, SAT, TAEs'. ' 

WARM). Bexar C( 


>?ers 14241 (BRIT/SMU). Burn 
.5427.5 (BRIT/SMU). Comal 

riLTofUS2<SlaiKlRRJl r 


Park, La Grange West Quad., in moist sandy loam soil over calcareous sandstone 
lawn, 15 Mar 1986, Carr 7169 and Kutac {TAES). Frio Co.: Rt. 1581 from Pearsall i 
Frio River crossing, dense herbaceous growth in shade of trees, loam soil, elev. ca. 
Mar \9^5,Ertter 3570 and Bear {CAS). Gillespie Co.: along stream in NE corner of 
of Coal Creek, common on N slope under oaks, 29 Apr 1959, Correll2l 1 68 andjohns 
SMU, MO, UC). Hayes Co.: San Marcos, dry rocky ground, 17 Apr 19\1 , Palmer 1 1 
Hill Co.: 7 mi WSW of Whitney, in detritus under limestone cliffs above dam, in tu: 



11 NE of Gra 

e slope above 

1 May 1949 J 

S. & G. Jones 7380 (BRIT/SMU, ccb, MICH, MO, SAT, TAES, TEX, US, VDB, WARM). 
Kerr Co.: 10.8 mi SW from Hunt on TX 39 from its jet. with FR 1340, mesic riverine habitat 
along W side ofthe South Fork of the Guadalupe River, 1 1 May 199 1, Jones 6670 and Krai 
(BRIT/SMU, ctb, MICH, MO, SAT, TAES, TEX, US, VDB, WARM). Kimble Co.: ca. 20 mi 
SW of Junction, valley of Paint Creek near its jet. with South Fork of Llano River, forest of large 
oaks, walnuts, and pecans, 1 1 May 1 947, McVaugb 8278 (CAS, NA, TEX). Lampasas Co. : 4. 1 
mi E on FM 580 from its jet. with FM 581, infrequent, 20 Apr 1988, S. & G.Jones 1268 
(TAES). Llano Co.: ca. 9 mi W of Buchanan Dam, on the road between Burnet and Llano, 29 
Apr 1946, Lundell 14363 (TEX). McClennan Co.: vicinity of China Spring, limestone hills, 
15 Apr 1970, Mauldtns.n. (BRIT/SMU). Mills Co.: 8 mi SW of Goldthwaite, creek bank, in 
shade, sand and limestone gravel, 30 Apr I960, Shinmrs 28312 (BRIT/SMU). Palo Pinto Co.: 
2.3 mi S on FR 4 from its jet. with FR 3137, S of Palo Pinto, mesic live oak W-facing hillside 
slope with reddish sandy loam soil and sandstone, 23 Apr 1992, 5. & G.Jones 8349 (Type series: 
BRIT/SMU, ctb, MICH, MO, SAT, TAES, TEX, US, VDB, WARM). Parker Co.: 9-5 mi E of 
Weatherford, clayey limestone soil near pond, in shade, 24 Apr 1949, Shinners 10963 (BRIT/ 
SMU). San Saba Co.: on Ellison Ranch, at Chapel, low, wet, shaded area by pool below natural 
waterfall, at edge of water, growing in black soil, 24 Apr 1977, Barnette236 (BRIT/SMU). 
Somervell Co.: 1 mi NE of Glen Rose on Hwy 67, sandy stream bank, in thicket, 2 Apr 1950, 
Shinners 12157 (BRIT/SMU). Sutton Co.: Sonora, experiment station, 21 Apr 1951 Jones 








—11 1 

|V JiiSSw 



1 Mi 






200 km \ ~ 


28383 (UC). Tarrant Co.: Sycamore Park, NW of East Rosedale St. and South Beech in Fort 
Worth, mesic creek meander through open woods with loamy clay soil, 23 Apr 1992, S. & G. 
Jones 8339 (BRIT/SMU, ctb, MICH, MO, SAT, TAES, TEX, US, VDB, WARM). Taylor Co.: 
Abilene State Park, in deciduous woods along Elm Creek, 28 May 1943, To/stead 7330 (OMA, 
UC). Travis Co.: 0.2 mi SW on the Andrandok Trail with its jet. with Loop 390 in Bull Creek 
Park, Austin, S side of Bull Creek on N-facing slope of a juniper-oak woodland, 7 Apr ] 990, S. 
& G. Jones 4223 (BRIT/SMU, ctb, MICH, MO, SAT, TAES, TEX, US, VDB, WARM). Uvalde 
Co.: near Uvalde, woods along Leona River, 30 Apr 1928, Palmer 33641 (UC). Williamson 

Whitehouse 18073 (BRIT/SMU). 

The following dichotomous key is for the taxa of section Phaestoglochin occur- 
ring in Arkansas, Louisiana, Oklahoma, and Texas. Incomplete veins refer to 
veins that do not extend from the base of the perigynium to the apex. Septate- 

nodules refer to cross-veins. Dorsal and ^ 
abaxial and adaxial respectively. 


Leafsheaths baggy around the culm (loose) 

2. Apex of ventral leaf sheath straight or slightly concave, not callused or only 
slightly thickened, friable; dorsal leaf sheath white or pale green with darker 
green veins with darker green septate-nodules, but not green, mottled with 
white, frequently with scattered red dots ventrally; perigynia turning stra- 

2. Apex of ventral leaf she: 

ith concave, callused or not, friable or not; some 

dorsal leafsheaths white 

or pale green with darke. 

: green veins and darker 

green septate-nodules wi 

th some dorsal sheaths green, mottled white, with- 

out scattered red dots ven 

traliy; perigynia remaining 

; green at maturity; peri- 

gynia 3.3-4.6 mm long. 

3. Spikes of inflorescence aggregated, apex of veni 

:ral leaf sheath concave. 


used, not friable; awns of m 

ost pistillate scales reach- 

,ng or exceeding the t 


ik; beaks 1. 0-1. 8(-2.0) 

mm long; widest leave 


wide; perigynia (3.4)3.6 

4.5 mm long 

3. Spikes of inflorescence 

? separated, internodesbei 

cween spikes frequently 

much greater than the length of the spikes; apt 

;x of ventral leaf sheath 

concave but not callu: 

than the base of the per 

igv'nium beak, beaks 8- 

L 0(-l 2)mmlong,wid- 

est leaves (4-)5-IOmr 


i-4 3) mm long C spargamoidt 

. Leafsheaths tight arotrnd tl- 

>e culm (not baggy) 

4. Lowest inflorescence brae 

t5 5-2=)cmlong,grcatlj 

' exceeding the mflores- 

cence, two to many time' 

.as long 

5. Culms smooth belo%. 

floodplain habitats 



5. Culms antrorsely scabc 

submesic woodlands 



4. Lowest inflorescence bract less than 5 5 cm long, no 

t exceeding rhc mflores- 

cence, or less than two tir 

Ties as long 

8. Perigynia ovate-lanceOid, veins present on ventral Sutiace, 
proximally over an enlarged spongy area at base of perigyni; 
9. Perigynia 1.3-1.8 mm wide; widest leaf blade 1 .5- 3.0 n- 

Perigynia spongy at base with or without a ventral swollen area 
11. Perigynia(1.4-)1.5-2.7(-2.8)mm wide, ovate-deltoid or con- 
base on ventral surface 


veins dorsally 

12. Perigynia (3. 3-)3.' 

veins dorsally 

11. Perigynia 0.9-1.8 mn- 
oblong; perigynia with 


13. Perigynia (3-)4-5 

4-5.2(-5.6) mm long; 0-5(-8) narrow 
Tim wide) ventrally; 0-10(-l 1) narrow 

a swollen spongy area at base of ventral 
times as long as wide; with rhizomes ± 


or only slightly 

14. Widest leaf wi 

times as long as wide; ^. 

'ithouc elongate 
stigmas once to 


■ (san'di3'ape"r-likeTexcep 


tTomedres°paringly Zng La^r veins 


. Apex of the ventral leaf sheath straight or slightly con- 
cave not callused or only slightly thickened, friable; fre- 
quently with scattered reddish dots; dorsal leaf sheath 
white or pale green with darker green veins with darker 
green septate-nodules, but not green, mottled with 

forming greater than 70° angle with the ground 

. Apex of ventral leaf sheath concave, callused, not fri- 
able; without scattered reddish dots; all dorsal leaf sheaths 

than a 50° angle with the ground, usually much 
less; many bract and/or pistillate scale awns greatly 

exceeding the perigynia beaks C.aintrina 

18. Some dorsal leaf sheaths white or pale green with 
darker green veins and darker green septate-nod- 
ules, but some sheaths green mottled with white; 
perigynia remaining green at maturity; widest leaves 
(3.0-)3.5-5.0(6.0) mm wide; most culms forming 
an angle greater than 70° with the ground; most 

apex of the perigynia beaks, occasionally a few bract 

awns surpass the beaks C.aggregata 

16. Perigynia 2.0-3.5 mm long, 1.3-2.3(-2.4)mm wide 19 

19- Perigynia bodies ovate-deltoid; perigynia beaks 0.3- 
0.7(-0.8) mm long with a single row of serrations, 
abruptly arising from rhe apex of the perigynium; wid- 

culm 2-6(-7); culm width, ca. 2 cm above rootstock, 
1.0-2.4(-3.5)mm wide; pistillate scale(1.3-)l. 5 2.2 
(-2.5) mm long; pistilate scale awn 0-0.8(-l .0) mm 

sheath mostly green, infrequently green mottled with 

white dots C. cephalophora 

Abaxial, adaxial, or both leaf surfaces minutely papillose (sand 

(-23.0) mm long; ventral surface of perigynia veinless, dorsal 
surface veinless or rarely with 1-4 incomplete narrow veins 

surface of perigynia 0-15-veined, dorsal surface 0-12-vein£ 

21. Ventral surface of perigynia with (5-)6-l 5 conspicuous 

broad veins (ca. 0.5 mm wide); dorsal surface with (0-) 

21. Ventral surface of perigynia with 0-6(-8) narrow veins 

;tillate scale; 

i 5.0-4 



mid-stripe 3-v 



>. forming an a 


• of 50" or 

;tillate scale; 





usually form 


an angle of 




Beaks of perigyi 

abruptly ai 









! mm long; 

if shea 

tths frequent 

)ts; plants WK 

Texas east 

: to G. 

eorgia and nc 


to Canada 


central Texas north to Oklahoma C. perdentata 

This treatment does not recognize varieties oi Car ex gravida. However, fur- 
ther research may prove that infraspecific taxa are warranted {i.e., C. gravida 
var. lunelliana (Mackenzie) F.J. Hermann], but until that time, I have opted for 
a more conservative approach. In floras where C. gravida, C. aggregata, and C. 
sparganioides occur together, they have classically been separated from other 
members of section Phaestoglochin by having baggy sheaths. However, I have 
encountered specimens of C. gravida and C. aggregata, both in the field and on 
herbarium sheets, that have tight leaf sheaths. It is possible that a single reces- 
sive gene is responsibile for tight leaf sheaths, or developmental through ontog- 
eny. Another plausible explanation provided by A. A. Reznicek [(MICH) pers. 
comm.} is that the individuals with tighter leaf sheaths are growing in less than 
optimum habitat. Regardless, C. gravida and C. aggregata usually have baggy 
leaf sheaths but can have tight leaf sheaths so they key out under baggy and 
tight leaf sheaths. 

The classical spelling of C muhlenbergii should be corrected to C. muehknhergii . 
This was pointed out by Peter Ball {(TRTE) pers. comm.}. Willdenow's original 
spelling was C. muhlenbergii with a diacritical sign (an umlaut over the "u"). 
Greuter et al. (1988), Article 73.6 state that diacritical signs are not used in 
Latin plant names. In names which are drawn from words in which such signs 
appear, the signs are to be suppressed with the necessary transcription of the 
letters so modified (i.e., u becomes ue). 

Carex perdentata s, closest putative relative is C. mesochorea. Carex perdentata 
differs in having longer perigynia with their bases slightly spongy, slightly nar- 

Jones, New species of Carex from Oklahoma and Texas 353 

rower and conspicuously longer leaves, usually longer inflorescences, more spikes 
per inflorescence but with fewer perigynia per spike, and for the most part, 
longer anthers. The teeth of the beaks of C. perdentata are noticeably longer and 
spreading at maturity in contrast to the short and usually straight teeth of C. 
mesochorea which rarely reach 1 mm long. The species epithet "perdentata" refers 
to the conspicuous teeth on the beaks of the perigynia. 

I am grateful to Gretchen D. Jones (USDA, AWPMRU), my wife, for 
viewing this manuscript, for her tireless enthusiastic help in the field, but mi 
important for her encouragement, understanding, and patience throughout r 
research. I thank A.A. Reznicek (MICH) and J.K. Wipfif (TAES) for their mar 
script review. I am grateful to Paul A. Fryxell and A.A. Reznicek for correct! 
my Latin description, to Keith Westover for providing the illustration, and 
Peter Ball (TRTE) for pointing out the orthographic correction of Ca. 

)uMORTiER, B.C. 1827. Florula Belgica, Operis Ui 

Casterman). P. 146. 
}reuter, W., et al. 1988. Chairman. International Code of Botanic 

by the Fourteenth International Botanical Congress. Koelt2 Scientific Books, Berlin. 
ajKENTHAL, G. 1909. Cyperaceae: Caricoideae. Das Pflanzenreich, IV, 20 (Heft 38):296-353. 

W. Engelmann, Leipzig, Germany. 
luNTH, C.S. 1837. Cyperographis synopticasiveEnumeratioCyperacearum. Vol. 2. Enumeratio 

Plantarum. StutgardiaeetTubingae. Pp. 378-381. 
/[ackenizie, K.K. 1931. North American Flora. 18. Cyperaceae, tribe 2, Carkae. New York 

Botanical Garden, Bronx, NY. 

E 1889. Cypera 

ceae Ii 

n: Engler und K. Prantl. Die naturlicl 

len Pflanzenft 

iglemann, Leipzig 


many. Pp. 98-130. 

.iCEK, A.A. and 1 

^.W. E 

mencan and Non 

[exico. Contr. Univ. Michigan Herb. lA 






Southern Weed Science Laboratory 
Stonmlle, MS 58776, U.S.A. 


Department of Biological Sciences 

Mississippi State University 

Mississippi State, MS 59762, U.S.A. 


Department of Biological Sciences 

Mississippi State University 

Mississippi State, MS 59762, U.S.A. 

do Carex godfreyi nuevo para A 
\. Se presentan extensiones de 
: indican la localidad y el habita 

In preparing a synoptic treatment oiCarex as a contribution to the Flora of 
Mississippi Project, the senior author has continued to examine herbarium speci- 
mens and conduct field surveys for species with potential to occur within JMis- 
sissippi and other southeastern states in the U.S.A. The authors have also con- 
tinued assessment of population size, distribution, and habitat requirements of 
recently reported species C. bicknellii Britton var. opaca F.J. Herm. and C. oklaho- 
mensis Mack., especially because these may have potential to become weedy. 
This mauscript adds to the knowledge oi Carex that has been reported in recent 
years (Bryson 1984a; Bryson& Jones 1990; Brysonet al. 1991; Brysonet al. 

1992; Bryson & Carter 1994; Carteret al. 1990; Morris & Bryson 1986; Naczi 
& Bryson 1 990). As previously discussed, the flora of Mississippi is still poorly 
known in comparison with several adjacent states (Bryson & Carter 1994). Lowe's 
Plants of Mississippi {\92\), although outdated, must continue to serve as a base- 
line for the general floristic work in the state. 

The terminology of physiographic regions or resource areas in Mississippi 
follows Lowe (1921) as adapted by Morris (1989). Herbarium abbreviations 
follow Holmgren et al. (1990), except ctb, MMNS, and USMH (pers. herb, of 
Charles T. Bryson; Mississippi Museum of Natural Science, Jackson; and Uni- 
versity of Southern Mississippi, Hattiesburg, respectively). 

Carex communis Bailey var. communis belongs to the section Acrocystis 
(Rettig 1988) and is known from dry to mesic rich woods in mountains, steep 
ravines, rocky ledges, and calcareous soils from Quebec to Ontario and Minne- 
sota, south to Georgia, Tennessee, Alabama, and Arkansas (Mackenzie 1931; 
Naczi 1993; Radford et al. 1964; Rettig 1988; Steyermark 1963). The follow- 
ing are the first collections of C communis from Mississippi. 

Voucher specimens: U.S.A. MISSISSIPPI. Monroe Co.: ca. 1 mi N Lake Monroe, Tl 3S R7E 
SIO SW/4, 12 May 1992, MacDonald4%l (ctb, IBE, MICH, others to be distributed); 28 Apr 

1993, MacDonald5928 & Warren (ctb, IBE, MICH, SWSL, others to be distributed); 19 May 

1994, Bryson 13651 & MacDonaU {cth , SWSL, others to be distributed). 

At this site in the Tennessee Hills Region, C. communis var. communis was 
found at an elevation of about 75 to 90 m in a mesic beech-maple-oak forest 
near the crest of a N- to NE-facing slope on Ruston and Cuthbert soils. It was 
associated with Acerbarbatum Mi&&., Aaaeapachypoda. Elliott, Adiantum pedatum 
'L.^Asimina triloba (L.) Dunal, Arabis canadensis L., Carex abscondita Mack., C. 
cephalophora Willd., C. corrugata Fernald, C. blanda Dewey, C. gracilescens Steudel, 
C. rosea Willd., C. laxtflora Lamarck var. serrulata F.J. Herm., C. willdemwii 
Wilid., Cynoglossum virginianum l..,Decumaria barbara L., Pagus grandifolia Ehrhart, 
Geranium maculatum \^.,Hepatica americana (DC.) Ker, Lindera benzoin (L.) Blume, 
Morus rubra L., Obolaria virginica L., Osmorhiza longistylis (Torrey) DC, Panax 
quinquefolium L., Quercus alba L., Rhamnus carolimana Walter, Thelypteris 
hexagonoptera (Michx.) Weath. 

Carex godfreyi Naczi occurs in wet hammocks, swamps, and floodplains in 
the coastal plain from southern North Carolina southward to the central penin- 
sula of Florida and west and to southwestern Georgia and nearby portions of the 
Florida panhandle. It inhabits shaded, mesic to wet areas in calcareous muck or 
sandy loam soils (Naczi 1993). Carex godfreyi belongs to a complex of species 
that include C. amphibola Steudel, C. corrugata Fernald, and C. grisea Wahlenb. 

Bryson, MacDonald and Warren, Notes on Carex 357 

Carex godfreyi is distinguished from the preceding species by its more loosely 
cespitose habit, its leaf sheaths and cataphylls with more extensive purple-red 
pigmentation, and its narrower leaf bases. The following citations are the first 
report of C. godfreyi from Alabama. 

Voucher specimens: U.S.A. ALABAMA. Houston Co.: ca. 9 mi S of Dothan, vicinity of Big 

(ctb, IBE, KNK, others to be distributed); Chattahoochee State Park, area where Irwin 
Mill Creek meets Hwy AL 95, 5 May 1993, MacDonald ^97 3 ( ctb, IBE, KNK, SWLS, 

At the site about nine miles south of Dothan, C. godfreyi grew in a Fagus 
grandifolia-Magnolia grandiflora L.-Magnolia virginiana L.-P inus glabra Walter- 
Quercus laurifolia y[\.z\\^.-Quercus michauxii Nutt. forest above Big Creek. Addi- 
tional associates included Carex abscondita, C. complanata Torr. & Hook., C. debiln 
iVLichx., C. leptalea Wahlenb., C. stylofkxa Buckley, Gordonia lasianthus (L.) El- 
liott, Uvulariafloridana Chapm. At the Chattahoochee State Park site, C. god- 
freyi grew in an Acer barbatum-Magnolia grandiflora-Magnolia virginiana-? ersea 
palustris (Raf.) ?) hemisphaerica Bartram forest above Irwin Mill Creek 
on a clay soil. Associates present at this site also included Aristolochta serpentaria 
L., Bumelia lanuginosa (Michx.) Persoon, Conopholis americana (L.) Wahl., Cornus 
stricta L., Dry op fens ludoviciana (Kunze) Small, Ilex cassine L., /. montana Torr. & 
Gray, Lyonia lucida (Lamarck) K. Koch, Osmanthus americana (L.) Benth. & Hook., 
Pedicularis canadensis L. , Ponthieva racemosa (Walter) C. Mohr, Rhododendron canescens 
(JVLichx.) Sweet, Sanicula canadensis L. , S. marilandica L. , Thelypteris palustris Schott, 
and Viburnum obovatum Walter. Carex atlantica Bailey subsp. capillacea (Bailey) 
Reznicek, C. leptalea, Ctcuta mexicana Coult. & Rose, Decodon verticillatus (L.) 
Elliott, Nasturtium microphyllum (Boenn.) Rchb., Rosa palustris Marshall, and 
Ztzamopsts miliacea (Michx.) Doell & Asch. grew in wetter areas nearby. 
Both collections are from the Doughtery Plain District of the East Gulf 
Coastal Plain of Alabama (Sapp & Emplaincourt 1975). 

Carex scoparia Willd. var. scoparia is known from open areas in swampy or 
wet river bottoms, valleys, prairie swales, upland prairies, margins of sink-hole 
ponds, and roadside ditches from Newfoundland to British Columbia, south to 
South Carolina, Tennessee, Arkansas, Oklahoma, New Mexico, and Oregon 
(Kolstad 1986; Mackenzie 1931; Radford et al. 1964; Steyermark 1963). This 
collection is first of C. scoparia from Mississippi. 

Voucher specimens: U.S.A. MISSISSIPPI. Lafayette Co.: Presbyterian Camp Hopewell, ca. 
6 mi NE Oxford, T8S R2W S9, 28 May 1994, Bryson 13910 (ctb, IBE, MICH, SWSL, others 

At this site in the Central Hills Region, C. scoparia var. scoparia grew in an 
open area on coarse sandy soil above a small spring-fed lake in association with 

Carex albolutescens Schwein., C.festucacea Willd., C. laevivaginata (Kiik.) Mack., 
C. longii Mack., C. lunda Wahlenb., C. triangularis Boeck., C. vulpinoidm Michx., 
Fuirena squarrosa Michx., Rhynchospora glomerata (L.) Vahl, Xyris tortaJ.E. Smith. 

Carex bicknellii Britton var. opaca RJ. Herm. was described from three 
collections of Delzie Demaree from river terraces in Lonoke and Prairie coun- 
ties, Arkansas (Hermann 1972). Carex bicknellii var. opaca was first found in 
Mississippi at a site in the Black Prairie Region of Mississippi where C. oklaho- 
mensis Mack, was first collected in the state (Bryson & Carter 1994). The follow- 
ing citation is the second report for Mississippi. 

Hwy US 78 and Dorsey- 

Three plants of C bicknellii var. opaca were observed at this site in close asso- 
ciation with C. oklahomensis Mack. Each plant was depauperate compared to 
those from a Lee County site observed in 1993 (Bryson & Carter 1994), but 
plant size was also reduced at the Lee County site in 1994, possibly due to dry 
conditions. In Itawamba County, C. bicknellii var. opaca was also associated with 
C. bmhii Mack., C. cherokeensis Schweinitz, C. complanata Torr. & Hook., C. glau- 
codea Tuck., C. longii Mack., C. vulpinoidm Michx., Cyperus echinatm (L.) Woods, 
C. lancastriensis Porter in Gray, C. odoratus L., C. strigosus L., Festucapratensis 
Huds., Eleocharis obtusa (Willd.) Schult., Fimbristylis autumnalis (L.) Roem. & 
Schult., and Rhynchospora capitellata (Michx.) Vahl. 

Carex oklahomensis Mack, is known in Mississippi only from Lee 
County, just east of Tupelo (Bryson et al. 1992). The following data are for 
an additional site in Lee County and for the first stations in Itawamba and 

Voucher specimens; U.S.A. MISSISSIPPI. Itawamba Co.: N of Dorsey, SW of jet. of Hwy 
US 78 and Dorsey-Fawn Grove Road Exit, T9S R7E S25, 24 May 1994, Bryson 13803 (ctb, 
MICH). Lee Co.: Tupelo, SW of jet. Hwy US 78 and Veterans Blvd. (=old Saltillo Road), T9S 
R6E S21, 24 May 1994, Bryson 13806 (ctb, MICH). Lowndes Co.: ca. 5 mi N of Columbus, 
Columbus Air Force Base, Tl6S R18W S30 SE/4 of E/2, 7 Jun 1993, Warren 2469 (ctb, IBE, 
MICH, MISS, MMNS, SWSL, USMH, VDB, VSC, additional specimens to be distributed); 19 
May 1994, Warren 2395, MacDonaid & Bryson (IBE, MICH, additional specimens to be distrib- 
uted); MacDona/d7197. Warren & Bfjson (ctb, IBE, MICH, additional specimens to be distrib- 
uted); Bryson 13680, Warren & MacDonald {^RYT ISmj , ctb, DSC, IBE, MICH, MISS, MMNS, 
USMH, VDB, VSC, additional specimens to be distributed). 

The additional collection from Lee County and the one from Itawamba County 
are both along highway US 78. As previously speculated (Bryson et al. 1992), 
it is likely C. oklahomensis was introduced with blown hay for erosion control 

Bryson, MacDonald and Warren, Notes on Carex 359 

along the construction corridor of Highway US 78 in Itawamba and Lee coun- 
ties. Associates at both sites are the same as those previously discussed for C. 
hicknellii var. opaca. It is likely that C. oklahommm was introduced at the Lowndes 
County site in much the same manner as the Itawamba and Lee county sites or 
as a contaminate of grass seeds planted for erosion control. At the Lowndes 
County site, C. oklahamensis grew west of Independence Boulevard along and 
south of a small drainage ditch through a pine-hardwood forest. In 1993, C. 
oklahomensis plants were 1.4 to 1.7 m tall. At the time the specimens were 
collected in June, plants were drooping from the weight of the infructescences. 
Plants were only 1.0 to 1.2 m tall in 1994; the reduced plant height may have 
been caused by increased competition from grasses and drier than normal con- 
ditions. Associates at the Lowndes County site are Carex frankii Kunth, C. 
longii Mack., C. triangularis Boeck., Eleocharis obtusa (Willd.) Schult., Festuca 
pratensis, and Glyceria septentrional is Hitch. In Mississippi, C. oklahomensis in- 
habits open sites with mucky clay soil, which are transitional between the 
Black Prairie and Tennessee Hills regions. 


Thanks are expressed to A.A. Reznicek (MICH) who confirmed the determi- 
nation oi Carex hicknellii var. opaca, C. oklahomensis and C. scoparia; R.F.C. Naczi 
(KNK) who confirmed the determinarions of Carexgodfreyi; and Richard Carter 
(VSC), S.D. Jones (TAES), R.F.C. Naczi, and A.A. Reznicek who reviewed the 

Bryson, C.T. and S.D.Jones. 1990. Carex comosa (Cyperaceae) new to Mississippi. Sic 

Bryson, C.T., S.W. Rosso, and R.F.C. Naczi. 1991. Carex baltzellti (Cyperaceae) ne? 

sippi with notes on Carex puta and Carex mpressinervia in Mississippi. Sida 14:4^ 

Bryson, C.T., R.F.C. Naczi, and S. McDaniel. 1992. Notes on noteworthy recor. 

Bryson, C.T. and R. Carter. 1994. Notes on Carex, Cyperus, and Kyllinga Cyperace 

Carter, R.M.W. Morris, and C.T. Bryson. 1990. Some rare or otherwise interest!] 

lants from the Delta Region of Mississippi. Castanea 55:40-55. 
Hermann, F.J. 1972. A new variety oiCarex hicknellii from Arkansas. Sida 5:49- 
Holmgren, RK., N.H. Holmgren, and L.C. BARNETT(eds.). 1990 Index herbarion 

KoiSTAD, A.O. 1986. Cyperaceae. In: Flora of the Great Plains. R.L. McGregor, T.I 

Morris, M.W. and C.T. Bryson. 1986. Carex swanii in Mississippi. Castanea 51:226-22 

Morris, M.W. 1989. Spiranthes (Orchidaceae) in Mississippi. Selbyana 1 1 :39^8. 

Naczi, R.F.C. and C.T. Bryson. 1990. Noteworthy records oi Carex (Cyperaceae) from tl 

southeastern United States. Bartonia 56:49-58. 
Naczi, R.F.C. 1993. Carex hrysonii and Carex godfreyi new species oiCarex section Gnseae froi 

the southeastern United States. Contr. Univ. Michigan Herb. 19:195-205. 
Radford, A.E., H.E. Ahles, and C.R. Bell. 1964. Manual of the vascular flora of the Carolina 

The University of North Carolina Press, Chapel Hill. 
Rettig, J.H. 1988. A biosystematic study of the Carex pensylvanica group (section Acrocystis) i 

North America. Ph.D. dissertation, University of Georgia, Athens. 
Sapp, CD. and J. Emplaincourt. 1975. Physiographic regions of Alabama. Geological Survc 


1963. Flora of Missouri. The Iowa State University Press, Ames. 



Botanical Research Institute of Texas 

^09 Pecan Street 

Fort Worth, TX 76102-4060, U.S.A. 

)oly is described and illustrated, and its phylogenetic relationship 
jfArdisia capuronii in subgenus Akosmos Mez is justified. A key ti 
1 Madagascar is presented. 

During a recent visit to the Laboratoire de Phanerogamie (P), to begin to 
prepare treatments of the Myrsinaceae for several different floristic projects, some 
curious collections by Capuron were discovered among the materials from Mada- 
gascar, representing a new taxon that is here described. 

Ardisia (subg. Akosmos) capuronii Pipoly, sp. nov. (Fig. 1) 

Quoad ramulos crassos, suberosos glabrosque, folia petiolata marginataque, inflorescentiam 
pyramidali-paniculatam longipedunculatamque, anteras ad initium poris apicalibus denique 

scrobiculatis (nee laevibus), petiolis 2-plo minoribus, pedunci 
epunctato (nee dense manifeste atro punctato-lineato), necnon florit 
dipositas atque lobulis calycinis rugosis (non laevibus) suborbicularibus (nee ovatis) praeclare 

Tree to 25 m tall; branchlets terete, 1.5-2 cm diam., glabrous, with exfoliat- 
ing reddish outer bark. Leaves persistent; lamina thinly coriaceous, obovate, 
(10-)12-20 cm long, (5-)(^10 cm wide, the apex widely rounded to truncate, 
the base cuneate, decurrent on the petiole, the midrib slightly raised above, 
prominently raised below, the secondary veins 14-18 pairs, densely and promi- 
nently black punctate, glabrous, the margin revolute; petiole marginate, 1.5-2 
cm long, glabrous. Inflorescences axillary, pendent, 21-35.8 cm long, 10-16 
cm wide, essentially columnar; inflorescence bract unknown; rachis hollow, gla- 
brous, densely and prominently black punctate; peduncle (7-)10-l6 cm long. 

PiPOLY, Ardisia in Madagascar 363 

broadened at base, glabrous; secondary branch bracts early caducous, unknown; 
floral bracts chartaceous, linear 2-3 mm long, 0.3-0.5 mm wide, the apex acumi- 
nate, densely and prominently red punctate, somewhat involute, the margin 
hyaline, entire, glabrous; pedicels cylindrical, recurved, 1 2-1 5 mm long, sparsely 
but prominently black and red punctate;. Flowers bisexual, pink, in terminal 
indeterminate 5-10-flowered umbels; buds ovoid, abruptly tapered apically; 
sepals 5, quincuncial, membranaceous, broadly ovate to suborbicular, 2.0-2.3 
mm long, 1 .8-2.0 mm wide, the apex broadly rounded, medially thickened or 
somewhat carinate, somewhat rugose and prominently black punctate basally, 
red punctate toward apex, the margin irregular, entire, hyaline, glabrous; petals 
5, quincuncial, membranaceous, ovate, 9-1 1 mm long, 3.5-4 mm wide, united 
ca. 1-1.5 mm basally, the apex long acuminate, slightly asymmetrical, sparsely 
but prominently black punctate, glabrous, the margin entire, opaque, glabrous; 
stamens free, subequalling petals, 8-10.5 mm long, the filaments free, flat, 
membranaceous, lanceloid, 2-2.5 mm long, glabrous, epunctate, the anthers 
linear, 6-7 mm long, 1 .8-2.5 mm wide, the apex acute, the base subsagittate, 
dehisceing first by apical pores, then by narrow longitudinal slits, the connec- 
tive densely and prominently black punctate-lineate dorsally; ovary ovoid, 1— 
1.5 mm long and in diam., black-punctate, placenta globose, ovules 3-4, unis- 
eriate; style tortuous, 7.5-8.5 mm long, densely black-punctate; stigma 
punctiform. Fruit unknown. 

Type. MADAGASCAR. Ambinanifaho: W of Isahana-Ambodipont, between Antalaha and 
Sambava; 19 Apr 1966 (fl), R. Capuron 24739-SF (holotype: P; isotypes: P- 2 sheets, TAN). 
' - -Ambohitralanana (Antalaha), 18 Mar 1967 


Local names: "Barabahala." 

Ecology and distribution: Ardisia capiironii is an element of the NE wet forest. It 
appears to be endemic to the region. 

Etymology: It is a great pleasure to name this new species for the late R. Capu- 
ron, collector and ardent student of the Malagasy flora. 

Ardisia capiironii is the fourth species oi Ardisia subgenus Akosmos Mez to be 
discovered in iVladagascar. It is closely related to A. procera Capuron, but is easily 

shorter petioles, stout peduncles, umbellate flowers, rugose and suborbicular 
calyx lobes. The four species oi Ardisia Sw. which occur in Madagascar all be- 
long to subgenus Akosmos (Mez, 1902) defined by paniculate, long-pedunculate 
inflorescences subequal to the leaves, slightly curved style subequal to the pet- 
als, punctiform stigma and anthers with first poricidal, then longitudinal dehis- 
cence. Miller and Pipoly (1 993) placed Ardisia subgenus Madardisia Capuron 
(1963) in synonymy under suhgenns. Akosmos, and the latter concluded that 
Stone ( 1 989, 1 990) was correct in his conclusion that Ardisia subgenus Akosmos 

rincipal center of diversity in the Malesian region. The species of An/i- 
ring in Madagascar are separated in the following key: 


a panicle of, 

umbels; panicle 


; perpendicular 

lis; placenta 

I rounded. 

. Branchlecs 

8-10 mm d 

lam.; leaf blades oblong or lanceolate, 2 

sepals char 

taceous, ova 

te, 1.8-2 mm, fl 


! chartaceous, o 

. Branchlets 

15-20 mm ( 

Jiam.; leaf blades obovate 

,5-10 cm wide 


5, broadly ov 

ate to suborbicL 

liar, 2-2. 

3 mm long, so 

nate; petal 

is membram 

iceous, ovate, 9- 

-11 mm 

long, slightly 

sparsely bt 


a panicle of I 




, Inflorescer 

mes; petioles ^ 

pedicels 1- 

-2 mm long 

; petals 4-5 mm 


, Inflorescer 

ice a pender 


corymbs; petic 

pedicels 10-20 mm long; petals 10-1 


ig, apices abn, 


I am grateful to Philippe Morat, Claude Sastre, Alicia Lourteig, Serge Bar- 
rier, Jean-Noel Labbat, and Odile Poncy, of the Laboratuire de Phaneroganne 
(P) for their hospitality during my visit, and to Pete Lowry (MO) and who 
kindly provided office facilities and logistical support. Linda Ellis executed the 
illustration with her usual accuracy and detail. 

Capuron, R. 1963. Contributions a I'etude de la flore de Madagascar. XIV. Le Genre Anlis/ 
Swartz (Myrsinacees) a Madagascar. Adansonia, ser. 2, 3:380-385. 

Miller, J. and J. Pipoly. 1993. A new species ofAn/isia from Madagascar. Novon 3:63-65. 
Stone, B.C. 1989- New and noteworthy Malesian Myrsinaceae, III. On the genus Ardhia Sw. 1 

Borneo. Proc. Acad. Nat. Sci. Philadelphia 141:263-306. 
1990. Studies in Malesian Myrsinaceae: 5. Additional new species of /\;r//.f/./Sv 

Proc. Acad. Nat. Sci. Philadelphia 142:21-58. 





Bee County College 

3800 Charco Road 

BeevilkTX 78102, U.S.A. 


Department of Biology 
Southwest Texas State University 
San Marcos, TX 78666, U.S.A. 

weed, are reported from the Comal and San 
nd illustrations of the species are provided. 

dc convertirse en una mala hierba acuatica, de los rios Comal y San Marcos de Tejas central, y se 

Key words: Hygrophila, Acanthaceae, aquatic weeds, Texas. 

In the spring of 1994 we began to question the identification of several aquatic 
macrophyte collections from the upper San Marcos River in Hays County, Texas, 
on deposit at SWT. The specimens in question had been variously identified as 
either Hygrophila lacustris (Schlecht. & Cham.) Nees (Acanthaceae) or Ludwigta 
repens Forst. (Onagraceae) and represented vouchers from two separate studies of 
the river's macrophyte flora (Lemke 1989, Staton 1992). Contributing to the 
difficulty of making an accurate determination was the fact that most of the 
specimens comprised only sterile material, a common deficiency of aquatic plant 
collections. Ultimately, however, we were able to secure both flowering and 
fruiting material of these plants and to identify them as Hygrophila poly sperma 
(Roxb.) T. Anderson, a previously unreported vascular hydrophyte from Texas 
that has the potential to become a troublesome aquatic weed. 

Hygrophila R.Br, comprises approximately 80 species distributed primarily 
in the Old World tropics, particularly Indochina and Malaysia, with only a few 
African and American species (Long 1970). The only representative of the ge- 
nus native to the U.S. is H. lacustris, which is distributed from Florida to eastern 

366 SiDA 16(2) 1994 

Texas (Correll & Correll 1975, Godfrey & Wooten 1981). Hygrophilapolyspernm 
is a native of India and Malaysia that was introduced into the United States in the 
1 940s and quickly became a popular aquarium plant (Innes 1 947). In the U.S. , 
the species has been reported as naturalized in lakes and drainage canals in south 
Florida (Les & Wunderlin 1981). Our recent field observations and collections 
indicate that the species is also naturalized in and along the San Marcos and 
Comal rivers in, respectively, Hays and Comal counties, Texas. Furthermore, 
herbarium records indicate that H. polyspemia has been well-established in the 
San Marcos River for at least 25 years (see specimen citations below). 

We surmise that H. polyspemia was introduced into Texas river systems either 
directly through cultivation by local aquatic plant nurseries, as documented by 
Hannan (1969) for the hydrophytic pteridophyte Ceratopteris thalictroides (L.) 
Brongn., or indirectly through careless dumping by aquarists. Profuse vegeta- 
tive reproduction is well developed m H. polysperma (Spencer & Bowes 1985, 
Van Dijk et al. 1986) and even small fragments will produce roots and grow 
into new individuals. We therefore feel it is likely that isolated introductions of 
plants cultivated for sale may have resulted in the establishment of the species 
in both the Comal and San Marcos river systems. 

The high growth potential of H. polysperma may pose a serious threat to the 
native flora and biotic integrity of the Comal and San Marcos river ecosystems. 
Several studies (Lemke 1989, Staton 1992, U.S. Fish and Wildlife Service 1994) 
have suggested that elements of the native biota of these two river systems are 
being displaced or otherwise adversely affected by exotic plant species. High 
growth potential, profuse vegetative reproduction, lack of seasonal variation in 
biomass, low light compensation and saturation points, a low CO2 compensa- 
tion point, and the capacity to rapidly change resource acquisition ability in re- 
sponse to environmental change are characteristics that make H. polysperma a 
competitive plant and potentially serious weed (Spencer & Bowes 1985, Botts 
et al. 1990, Kovach et al. 1992). The species is included on the federal list of 
noxious aquatic weeds (U.S. Department of Agriculture 1983) and is listed as a 
category II species by the Exotic Pest Plant Council of the State of Florida, in- 
dicating that its population is rapidly expanding and has the potential to invade 
and disrupt native vegetation in that state (Lantz 1993); however, the species is 
not currently recognized as a potentially harmful aquatic weed in Texas. 

To facilitate the identification of future collections of this species, we provide 
the following key, description and illustration (Fig. 1): 


ND Lemke, Hygrophila polysperma from Texas 

Fig 1 Hygrophila polysperma, A Distal p 
roots B Terrestrial form, drawn to sam 

368 SiDA 16(2) 1994 

Hygrophilapolysperma(Roxb.)T. Anderson, J. Linn. Soc, Bot. 9:426. 1876. 

Justidapolysperma^oxh.,Y\.lnA. 1:120. 1820. 

Hemuk/phispo/ysperma {Roxh.)Nees in Wall., Pi. Asiat. Rar. 3:30. 1832. 

Perennial rhizomatous terrestrial or aquatic herbs to 1 .5 m tall. Stems ascen- 
dant or rarely erect, more or less 4-angled, puberulent to giabrate, with abun- 
dant elongate or rarely rounded cystoliths in the epidermis. Leaves opposite, 
broadly elliptic to oblanceolate, acute at apex, attenuate to a subpetiolar base, 
minutely denticulate to entire, 7-65 mm long, 2-10 mm wide, mostly gla- 
brous but those subtending the flowers hispid, especially on the margins, the 
abaxial and adaxial surfaces with abundant elongate cystoliths. Flowers solitary 
in the axils of uppermost leaves, sessile. Bracts narrowly lanceolate, 4-5 mm 
long, herbaceous. Calyx equally 5-lobed, the lobes scarious-margined, 4-5 mm 
long, united basally, hispid. Corolla bluish-white (yellowish in dried specimens), 
5-6 mm long, the upper lip 2-lobed, the lower lip 3-lobed, puberulent. Fertile 
stamens 2, included, filaments glabrous, anthers 2-celled, ca. 1 mm long. Ovary 
hispid distally, the style ca. 3 mm long, sparingly hispid, the stigma flattened, 
ca. 0.3 mm long. Capsule 6-7 mm long, mostly glabrous but with a few distal 
hairs. Seeds flattened, round, ca. 0.8 mm diameter. Blooming mostly Sep-Oct. 

In the San Marcos and Comal rivers, H. polysperma is most often found grow- 
ing completely submersed, although we have frequently encountered both emer- 
gent and terrestrial individuals along the upper San Marcos River. Submersed 
individuals have longer stems (0.3-1.5 m tall) with elongate internodes (20-54 
mm long), relatively large oblanceolate leaf blades (17-65 mm long), and pro- 
duce abundant adventitious roots at the upper nodes. Terrestrial individuals are 
of shorter stature (10-20 cm tall) with shorter internodes (2-16 mm long), 
smaller elliptic leaves (7-25 mm long), and bear roots almost exclusively on the 
rhizome. We have observed flowers and fruits primarily on terrestrial individu- 
als and, occasionally, on emergent shoots of partially submersed plants; in both 
cases the distal leaves subtending the flowers are marginally hispid. These ob- 
servations accord well with those of Sculthorpe (1967) who cited numerous 
examples of normally terrestrial plants (e.g., species oiAlisma L.,Bacopa Aublet, 
Campanula L., Gratiola L., Nomaphila BL, Ranunculus L., and Rotala L.) that can 
grow entirely submerged even in deep water, leading to their common use as 
ornamental plants in aquaria; submersed individuals of such species were usu- 
ally found to be characterized by elongation of the leaf blade, loss of pubescence. 

Two previous studies of the aquatic macrophytes of the San Marcos River 
failed to document the occurrence there of H. polysperma. Lemke (1989) incor- 
rectly identified collections of this species as H. lacustris, while Staton (1992) 
misidentified her collections as Ludivigia repens. Hygrophila lacustris, a native of 
the southeastern U.S., occurs in only a few counties in southeast Texas (Brazoria, 

Angerstein and Lemke, Hygrophila polysperma from Texas 369 

Chambers, Fort Bend, Hardin, Harris, Jackson, Montgomery, Orange, and 
Walker counties), where it grows as a terrestrial or emergent plant along muddy 
stream and pond margins (Wasshausen 1966, Correll & Correll 1975). The 
plants are typically erect herbs to 80 cm tall with leaves mostly 5-12 cm long 
and flowers borne in distinct axillary clusters along the length of the stem. 
Hygrophila polysperma, in contrast, is usually found submersed, the leaves of the 
aerial shoots are typically 7-25 mm long, and the flowers are solitary in the axils 
of the uppermost leaves. 

When first introduced to the aquarium plant market, H. polysperma was though 
to be a species oiLuckvigia L. and was given the common name "oriental ludwigia" 
(Innes 1 947). Vegetatively, the terrestrial shoots of H. polysperma are very similar 
to those of L. repens, a native species also known from the San Marcos and Comal 
, elliptic leaves and short internodes. 
jctures, however, the two species are 
readily separable by nodal morphology. The petioles of L. repens are subtended 
by a pair of minute, glandular stipules less than 1 mm long, while the connate 
leaf bases of H. polysperma are exstipulate but bear a number of setiform hairs to 
1.5 mm long (Fig. 2). 

Hygrophila polysperma is the second adventive aquatic member of the Acan- 
thaceae to be reported from Texas in recent years. Ramamoorthy and Turner 
( 1 992) documented the occurrence oiNomaphila stricta (Vahl) Nees, another 
Malaysian species, from San Felipe Springs in Val Verde County and surmised 
that this species was also introduced by aquarists. 



A ' - 

B p^' 

Fic. 2. Comparison of nodal morphology i 
B. Uuhcgia repens. Drawn by Amy L. Mai 


■ifu,. A. H,grol,h,l„l«l,.,pm,,.,. 

370 SiDA 16(2) 1994 

Specimens examined. TEXAS. Comal Co.: Comal River, Landa Park, City of New Braunfels, 
ca. 100 m upstream of Pecan Island, mid-channel, in 1-2 ft of water, 21 Feb 1994, Angerstein 
94-1 (BRIT, SWT, TEX); Comal River, Landa Park, City of New Braunfels, in spring run 
underneath Landa Dr., 23 Feb 1994, Angerstein 94-2 (SWT). Fiays Co.: San Marcos River 
below Southwest Texas State University campus, 30 Nov 1969, Tablers.n. (SWT); shoreline 
sandbar formed at mouth of Sessoms Creek, 22 Nov \91'b, Litchfield s.n. (SWT); San Marcos 
Riverdownstream from Clear Springs Aprs, spillway, 5 Jul \9lG, Litchfield s.n. (SWT); South- 
west Texas State University raceway in gravel substrate, 28 Aug 1 976, Litchfield s.n. (SWT); San 
Marcos River at Thompson's Island below county road bridge, 6 Mar 1 99 1 , Staton s. n. (SWT); 
San Marcos River upstream from Purgatory Island, 26 Apr 1991 , Staton s.n. (SWT); San Marcos 
River at University Blvd., 26jun \99\,Bierner91-7 1 (TEX); Rio Vista Park, Purgatory Is- 
land, City of San Marcos, 30 Mar 1994, Angeistein 94-5 (SWT, TAES); San Marcos River, Lions' 
Park area, San Marcos, 30 Mar 1994, Angerstein 94-6 (SWT); San Marcos River, Sewall Park, 
San Marcos, along edge of concrete channel, 30 Mar 1994, Angerstein 94-7 (SWT, TEX); South- 
west Texas State University campus pond beside Freeman Bldg., 30 Mar 1994, Angerstein 94-8 
(SWT, TAES); Southwest Texas State University campus pond directly in front of J.C. Kellam 
Bldg., 30 Mar 1994, Angerstein 94-9 (BRIT, SWT). 


We would like to thank the Parks and Recreation Department of the City of 
New Braunfels, Texas, for permission to collect in Landa Lake, the Center for 

Bot. 36:207-214. 
CoRRELL, D.S. and H.B. Correi 

States, 2 vols. Stanford Univ 
Godfrey, R.K. and J.W. Woot 

Innes, WT 1947. Hygrophila. a new aquarium plant. Aquarium (Brooklyn) 16:30-31 . 

KovACH, C.W, J.P KuRDZiBi,, R. Bowman, J. Wagner and J.M. Lawrence. 1992. The effect; 
of stress and disturbance on proximate composition, allocation of production, photosynthe- 
sis, respiration, and chlorophyll levels in Hygrophila polysperma (Roxb.) Anders. (Acanthaceae) 
Environm. Exp. Bot. 32:479-486. 

Lantz, PS. 1993. Florida's most invasive species. Palmetto 1 3:6-7. 

Lemke, D.E. 1989. Aquatic macrophytes of the upper San Marcos River, Flays Co., Texas 
Southw. Naturalist 32:289-291 . 

Les, D.H. and R.P Wunderlin. 19^1 . Hygrophila polysperma 
Sci. 44:189-192. 

Long, R.W. 1970. The genera of Acanthaceae in the south 
Arbor 51:257-309. 

AND Lemke, Hygrophila polysperma from T 
HY, T.P. and B.L. Turner. 1 992. Nomaphila s 

Spencer, W. and G. Bowes. 1985. Li 

San Marcos River. M.S. thesis, Southwest Texas State University, San Marcos. 
United States Department OF Agriculture. 19<S3. Noxious weeds. Fed. Reg. 48:20037-20047. 
United State.s Fish and Wildlife Service. 1994. Draft San Marcos and Comal Springs and 

associated aquatic ecosystems (revised) recovery plan. Albuquerque, NM. 
Van Dijk, G.M., D.D. Thayer and W.T Haller. 1986. Growth oiHygrophi/a and Hydrilla in 

flowing water. J. Aquatic PI. Managem. 24:85-87. 
Wasshausen, D.C. 1966. Acanthaceae. In: C.L. Lundell, ed. Flora of Texas 1:223-282. 



Department of Botany 

University of Texas at Austin 

Austin, TX 787 13-7640, U.S.A. 


Department of Entomology 

Texas A&M University 

College Station, TX 77845-247 5, U.S.A. 


Director of the Library 

New York Botanical Garden 

Bronx, NY 10438-3126, U.S.A. 

T.S. Brandegee and his wife Katharine Brandegee were important California 
botanists (Setchell 1 926) who contributed greatly to our knowledge of the flora 
of that floristically rich region during the early decades of the 20th century. The 
nature and extent of their contributions are amply dealt with elsewhere (see 
references) and need not detain us here. We became aware of the fact, however, 
that T.S. Brandegee 's full name is referred to in the literature with several vari- 
ants, which may be a source of confusion. We wished to ascertain which alterna- 
tive is correct and to share our findings. 

In his own publications Brandegee usually cited his name with initials only, 
as "T.S. Brandegee." Others often referred to him in this same way, but some- 
times also as "Townsend Stith Brandegee" (Lanjouw & Stafleu 1954, Lindsay 
1955, Thomas 1969, Stafleu & Cowan 1976), sometimes as "Townshend Stith 
Brandegee" (Setchell 1926, Jones 1929, Ewan 1950, Lindsay 1955, McVaugh 
1956,Langman 1964, Thomas 1969, Hunt Institute 1972, Lenz 1986, pp. 26, 
84, Bonta 1991, p. 87, Reveal & Pringle 1993), and sometimes as "Thomas 
Stith Brandegee." Qones 1929, Rodgers 1968, Lenz 1986, p. 200). Indeed, four 
of the references cited above Qones 1929, Lindsay 1955, Thomas 1969, Lenz 
1 986) are internally inconsistent in citing two different variants of Brandegee s 
name. We therefore conclude that no single reference can be considered authori- 
tative in solving this problem, nor do we believe that this sort of question can be 
resolved by majority rule. 

All references agree on his second name "Stith," taken from his mother's 

maiden name, so that is not a source of controversy. The first name (whether 
Thomas, Townsend, or Townshend), however, is clearly a source of uncertainty. 
But two references give an unequivocal answer and thus resolve the uncertainty, 
and we wish to share this information in the present note. 

One paper published by Brandegee (Brandegee 1913) departs from his usual 
practice of using only initials in the byline, and clearly states "Townshend Stith 
Brandegee." The second and more persuasive bit of evidence is the portrait of 
Brandegee at the age of 80 published by Setchell (1926: plate 14), which in- 
cludes the full signature of Brandegee, reading "Townshend Stith Brandegee." 
Thus, since it is attested in his own hand, there can be no doubt about the 
correct spelling of Brandegee 's given name, which was not "Townsend" and 
certainly was not "Thomas." 

Even more compelling but less readily accessible are documents in the Brande- 
gee archives kept at the herbarium of the University of California (UC), includ- 
ing his "doctoral diploma from Yale, pension papers, and discharge from the 
U.S. Army," all of which specify his given name as "Townshend" (Barbara Ertter, 
pers. comm.). 

BoNTA, M.M. 1991. Kate Brandegee, accidental botanist. In: Women in the Field. Texas Ai 

University Press. 1991. pp. 85-92. 
Brandegee, T.S. 1913. Plantae Mexicanae Purpusianae, V. Univ. Calif. Publ. Boc. 4:375-3 
EwAN, J. 1950. Rocky Mountain naturalists. University of Denver Press. 
Hunt Botanical Library. 1972. Biographical dictionary of botanists represented in the Hi 

Institute portrait collections. G.K. Hall & Co.: Boston. 
Jones, M.E. 1929. The Brandegees. Contr. W. Bot. 15:15-18. 

University ofPennsylvania Press. 1013 pp. 
Lanjouw, J. and F.A. Stafleu. 1954. Index herbariorum, Part II, Collectors (A-D). 
Lenz, L.W. 1986. Marcus E. Jones. Rancho Santa Ana Botanic Garden. 
Lindsay, G. 1955. Notes concerning the botanical explorers and exploration of Lower Calil 

nia, Mexico. Belvedere Scientific Fund. 
McVaugh, R. 1956. Edward Palmer, explorer of the American West. University of Oklaho 

Reveal, J.L. andJ.S. Pringle. 1993. Taxonomic botany and floristics. In: Flora of North Ame: 
Editorial Committee, eds. 1993 + . Flora of North America North of Mexico. 2+ vols. Oxf 
University Press, New York. Vol. l,pp. 157-192. 

York. ' 
Setchell, W.A. 1926. Townshend Stkh Brandegee and Mary Katharine (Layne) (Curran) Bran 

gee. Umv. Calif. Pubi. Bot. 13:155-178 + 2 plates. 
Stafleu, F.A. and R.S. Cowan. 1976. Taxonomic literature-II. Vol. 1 (A-G):303. 

GENTIANACEAE — New distribution records, based on specimens I have re- 
cently identified, are reported here for six Mexican and South American species 

Gentiana mirandae Para 

MEXICO. Veracruz: Mun. Huayacocotla, Viborillas, Huayacocotk, 2200 m, Hernandez M. 

This is the first record of this showy but rarely collected species for Veracruz, 
although the locality is near a previously reported locality in Hidalgo (Pringle 

Gentianella achalensis (Hieron. ex Gilg) TN. Ho & S. W Liu 

BOLIVIA. Tarija. Prov. Arce: 12.5 km N of Emboroza-Sidras road, on road to Padcaya, 

22° 12'S, 64° 37'W, 1150 m, Solomon 10160 (MO). Prov. Cercado: 54.9 km E of Tarija- 

Padcaya road, on road to Entre Rios, 21° 29'S, 64° 20'W, 2050 m, Solomon 10324 (MO). 
These records extend the known range of this species hitherto recorded only 

form Argentina, into southernmost Bolivia. Field work in Bolivia by J.C. Solomon 

was supported by the National Science Foundation. 

des (Griseb. ex Gilg) J.S. Pringle 
/. Loayza: along the S fork of the Rio Jij 

77320 (MO). 

This record extends the known range of this species, hitherto recorded only 
form Depto. Puno, Peru, into adjacent Bolivia. 

Gentianella primuloides (Gilg) J.S. Pringle 

BOLIVIA. La Paz. Prov. B. Saavedra: cerca de la carretera nueva de Cotacampa a Ulla- 
Ulla, Cerro Pifiita, 4500 m , Menhofer X- 1 782 (HAM, LPB). 

The isotype at G, from Depto. Puno, Peru, which is of particularly good 
quality, is closely matched by this recent collection from just across the border 
in Bolivia, indication that this species is correctly attributed to the floras of 
both countries. The corollas were described by Manhofer (in sched.) as red 
('rojas'), and show contrasting whitish bases, corresponding to Gilg s (1916) 

Gentianella vargasii Fabris 

BOLIVIA. La Paz. Prov. MuriUo: La Paz-Calacoto 64 km hacia el nevado Illimani, sobre el 
pueblo de Pinaya, pie de Illimani, 4200 m, Beck 9073 (HAM, LPB). 

This record extends the known range ca. 570 km to the southeast and is the 
first for BoHvia, the species hitherto being known only form Depto. Cuzco, Peru 
(Fabris 1958). This indicates that the range of G. vargasii may be more extensive 
than that of most Andean Gentianella species, even though the species may be rela- 
tively uncommon. A parallel is approached, however, in the relatively extensive 
range oiG.punicea (Wedd.) Holub, which extends from Depto. La Paz, Bolivia, 
along the mountain ranges northeast of Lake Titicaca, well into Peru. These 
ranges are essentially continuous from the international border to the southeast- 
ern part of Depto. Cuzco from which G. vargasii was previously reported. 

Tapeinostemon zamoranum Steyerm. 

PERU. Amazonas. Luya Prov.: Camporredondo-Tullanya, trocha hacia el Cerro Huicsocunga, 
2350 m, Diaz & Campos 5733 (HAM, MO). Cajamarca. Cutervo Prov.: San Andres de Cutervo, 
Parque Nacional de Cutervo, entrando por "Chorro Blanco" sobre invernas del Sr. Nicolas 
Navarro, 2300-2400 m, Diazetal. 3951 (HAM, MO). 

These records, which are the basis of my inclusion of this species in Brako & 
Zarucchi's ( 1 993) Catalogue of the Flowering Plants and Gymnosperms of Peru, are 
the first for the genus and the species for Peru, T. zamoranum hitherto having 
been reported only from southern Ecuador (Steyermark 195 1). 
— -JamesS. Prmgle, Royal Botanical Gardens, Box 399, Hamilton, Ontario, Canada 

This is Contribution No. 85 from the Royal Botanical Gardens, Hamilton, 

Brako, L., and J.L. Zarucchi. 1993. Catalogue of the flowering plants and Gymnosperms o) 
Peru/Catalogo de las Angiospermas y Gimnospermas de Peru. Monogr. Syst. Bot. Missour: 
Bot. Gard. 45 xl + 1286 pp. 

Fabris, H.A. 1958. Notas sobre Gentianella del Peru. Bol. Soc. Argent. Bot. 7:86-93. 

GiLG,E. 1916. GentianaceaeAndinae. Bot. Jahrb. Syst. 54(Beibl. 1I8):4-122. 

Macbride, J.F. 1959. Gentianaceae. Gentian Family. In: Flora of Peru. Field Mus. Nat. Hist. 
Bot.Ser. 13(5):270-363. 

Pringle, J.S. 1977. Taxonomy and distribution oiGentiana (Gentianaceae) in Mexico and Cen- 
tral America. I. Sect. Pneumonanthe. Sida 7:174-217. 

Steyermark, J.A. 1951. The genus Tapeinostemon (Gentianaceae), Lloydia 14:58-64 

ILLINOIS — Sagina procumbens L., procumbent pearlwort, is a Eurasian native 
that has been found at various sites in North America, well established prima- 
rily along both the Atlantic and Pacific coasts (Crow 1978). Within the Great 
Lakes Region, it has been reported fi-om scattered sites in iVLichigan, Minnesota, 
Ohio, and Wisconsin; all but four sites, three in the Lower Peninsula of Michi- 
gan (Crow 1978, Voss 1985) and one m central Ohio (Crow 1978), are along the 
shore of Lake Superior. 

Sagina procumbens can now be reported from four counties in Illinois from the 
following collections: 

ILLINOIS. Cook Co.: in patio blocks, 2601 Lake Ave., Wilmette, last week Jun 1976, E. 
McArdle&R. H. Fargo s.n. (MOR, SIU); sidewalk cracks, 5300 Block of N Spaulding Ave., 
Chicago, 27 Jun 1985, HaUaway3326 (MOR, [SIU]). Kane Co.: sidewalk cracks, 40 W 693 
McDonald Rd., N ofWasco, 13 Jul 1991, K. Johnson s.n. (MOR). Lake Co.: between stones or 
brick, Farwell & Lydia Lees residence, Lake Forest, 6[?} Jul 1965, i?a/«OK 49-X (SIU). Peoria 
Co.: in brick sidewalk cracks, 200 block N Garfield St., Peoria, 15 Jun 1955, Chase 14232 

The reports of Sagina decumhem (Elliott) Torrey & A. Gray from Cook and 
Lake counties by Mohlenbrock and Ladd (1978) and Swink and Wilhelm (1979) 
are based on specimens of 5. procumbens cited here. Sagina decumbens is native to 
sandy areas in the southeastern United States and, except for a single collection 
from Peoria Co., is "occasional in the s. 1/2 of the state (Mohlenbrock 1986)." 
When the dots for Cook and Lake counties are removed, the map in Mohlen- 
brock and Ladd (1978) for the distribution of 5. decumbens in Illinois is once 
again correct. 

Hattaway 3326 is the basis of his report of Arenaria groenlandica (Retz.) Sprengel 
as new to Illinois and the Midwest (Hattaway, 1987); it was erroneously identi- 
fied as that species. 

Sagina procumbens can be distinguished from S. decumbens via the following 

1. Plant perennial, stems procumbent, sepals diverging from the dehiscent 

1 . Plant annual, stems filiform and mostly erect, sepals remaining appressed to the 

dehiscent capsule S. decumbens 

Two other characters are commonly cited as separating these two species, 
features that are diagnostic in many, but not all cases. The flowers of S . procumbens 
are predominantly 4-merous with petals much shorter than the sepals or absent 
while those ofS. decumbens are predominantly 5-merous with petals about equal- 
ling the sepals. Some populations of 5^. decumbens include plants that are smaller 
and more branched and that often have 4-merous apetalous flowers (specimens 

SiDA 16(2) 1994 

seen from Coles and Wabash cos., ILLS); Crow (1978) included rhese v 
range of variation ofS. decumbens. 

— RichardK. Rabeler, University of Michigan Herbarium, North University Build- 
ing, Ann Arbor, Ml 48109-1037, U.S.A. 

Crow, G.E. 1978. A taxonomic revision of 5tf^/W (Caryophyllaceae) in Norcli America. Rhoclora 

Hattaway, R.A. 1987. Arenaria groenlandica (Retz.) Sprang, var. groenlandka in the Midwest. 

MOHLENBROCK, R.H. 1986. Guide to the vascular flora of Illinois, rev. ed. Southern Illinois 

Univ. Press, Carbondale & EdwardsviUe. 
MoHLENBROCK, R.H. and D.M. Ladd. 1978. Distribution of Illinois vascular plants. Southern 

Illinois Univ. Press, Carbondale & EdwardsviUe. 
SwiNK, F. and G. Wilhelm. 1979. Plants of the Chicago region, rev. ed. The Morton Arbore- 

DUCTION TO TEXAS— Emilia is an Old World genus with approximately 
45 species, three of which have become neotropical weeds (Nicolson 1975). 
Until recently Emilia fosbergii Nicolson has been reported in the United States 
only as a casual weed growing in southern Florida (Cronquist 1980; Barkley & 
Cronquist 1978). 

In the fall of 1993, a population of approximately 40 individuals of Emilia 
fosbergii was found growing in and around a gravelly path in a south Austin 
nursery. Upon inquiry the manager informed me that the nursery has suppliers 
in Florida, which might explain the presence of this weed. 

Voucher specimen: TEXAS. Travis Co.: City of Austin, A-1 Grass Nursery, two blocks S of 
Barton Skyway along S Lamar St., growing in gravel of parking lot and planting area, 20 Nov 

Emilia fosbergii is distinguished from other Texas genera in the Senecioneae 
by the red disk flowers and absent ray flowers. The receptacle is flat to slightly 
convex, the leaves are alternate. 

, Unwmity of Texas, Austin, TX78713, 

Barkley,T.M. and A. Cronquist. 1978. Emiiia. In: N. Amer. Fl., Ser. II, 10:147-150. 
Cronquist, a. 1980. Vascular flora of the southeastern United States. Vol. I. Asteraceae. Univ. 

North Carolina Press, Chapel Hill. 
N1COI.SON, D.H. 1975. Emilia fosbergii, a new species. Phytologia 32:33. 

spring systems that arise along the Balcones fault zone of central Texas support 
a diversity of aquatic macrophytes, including a number of adventive species that 
have not been reported from elsewhere in the state (Lemke 1989, Ramamoorthy 
& Turner 1992, Angerstein & Lemke 1994). The upper San Marcos River in 
Hays County, Texas, supports a macrophyte community comprising thirty-one 
species (Lemke 1989), two of which, Ceratoptens thalktroides (L.) Brongn. and 
Linmophila sessiltflora BL, have not previously been reported elsewhere in Texas 
(Correll & Johnston 1970). Recent collections of aquatic macrophytes fromLanda 
Lake, a small reservoir formed by the damming of the headwaters of the Comal 
River in New Braunfels, Comal County, Texas, have documented the presence 
of these two species in this river system as well. 

Ceratopteris thalktroides is a tropical, free-floating, homosporous aquatic fern 
that has been introduced into Florida, Louisiana, Texas, and California (Lloyd 
1993). The species was first reported from Texas by Morton (1967) and its 
introduction into the San Marcos River by a local aquarium plant supply com- 
pany was documented by Hannan (1969). The following collection represents 
only the second county record for the species in Texas: 

Voucher specimen: TEXAS. Comal Co.: free-floaring along north shore of Landa Lake, Landa 
Park, City of New Braunfels, l6Apr \99A,Umke4163 (SWT). 

Linmophila sessiliflora is a submersed or emergent macrophyte indigenous to 
India and Southeast Asia. It resembles the native species Cabomba cawliniana A. 
Gray in gross morphology, but can be easily distinguished in the vegetative 
state by its bright green coloration, more compact growth habit, and verticillate 
leaves. Limnophila sessiliflora is reported to be sporadically naturalized in Florida 
and Georgia; the following collection represents only the second county record 
for the species in Texas: 

Voucher specimen: Texas. Comal Co.: submerged along north shore of Landa Lake, Landa 

It seems likely that these species were introduced into Landa Lake either 
intentionally, as has been documented for C. thalictroides in Texas by Hannan 
(1969) and for L. sessiliflora in Fforida by Mahler (1980), or unintentionally 
through careless dumping by aquarists, as has been surmised for the aquatic 
weeds Nomaphila strkta fVahl) Nees (Acanthaceae) by Ramamoorthy & Turner 
(1992) and Hygrophilapolysperma (Roxb.) T Anders. (Acanthaceae) by Angerstein 

Although C. thalictroides has not been reported to occur so abundantly as to 
cause problems in aquatic ecosystems, L. sessiliflora is of concern as a potential 
aquatic weed. Several species oiLimnophila, including L. sessiliflora, are respon- 
sible for major weed infestations throughout much of Southeast Asia and virtu- 
ally all herbicides registered for use in aquatic systems have proven ineffective in 
controlling these species (Misra & Tripathy 1 97 5 , Takematsu et al. 1976, Mahler 
1980). Spencer & Bowes (1985) reported that L, sessiliflora has several character- 
istics that could provide it with a competitive advantage over native aquatic 
plants in Florida, such as a substantial reproductive capacity, the potential for a 
low photorespiration rate, and the ability to photosynthesize effectively under 
low light regimes. 

Ltmnophtla sessiliflora was reported to be uncommon in the San Marcos River 
by Lemke (1989) but has been found to be abundant in parts of Landa Lake, 
where several individuals were observed to be flowering in the summer of 1994. 
Although the species has a limited capacity for asexual reproduction, this may 
be offset by its profuse sexual reproduction. Spencer & Bowes (1985) reported 
that each flower of L. sessiliflora may set between 200 and 300 seeds with a 
germination rate as high as 96 percent. Presently there is no evidence that L. 
sessiliflora is having a deleterious effect on the growth of native aquatic plant 
species in either the San Marcos or Comal rivers; however, the spread of this 
species in the spring systems of central Texas should be closely monitored. 
— David E. Lemke, Department of Biology, Southwest Texas State University, San 
Marcos, TX 78666 U.S.A. 

Angerstein, M.B. and D.E. Lemke, 1994. First records of the aquatic weed Hygrophilapolysperma 
(Acanthaceae) from Texas. Sida 16:365-371. 

CoRRELL, D.S. and M.C. Johnston. 1970. Manual of the vascular plants of Texas. Texas Re- 
search Foundation, Rennet, TX. 

Hannan, H.H. 1969. The introduction and establishment oi Ceratoptens in Texas. Amer. Fern 
J. 59:122. 

Lemke, D.E. 1989. Aquatic macrophytes of the upper San Marcos River, Hays County, Texas. 

Lloyd, R.M. 1993. Parkenaceae. In: Editorial Committee, eds. Flora of North Amen 

of Mexico, vol. 2, pp. 1 19-121. Oxford Univ. Press, New York., M.J. 1980. Ltmnophila, a new exotic pest. Aquatics 2:4-7. 
MiSRA, G. and G. Tripathy. 1975. Studies on the control of aquatic weeds of Orissa 

Effect of chemical herbicides on some aquatic weeds. J. Indian Bot. Soc. 54:65-71 
Ramamoorthy, T.P and B.L. Turner. 1992. Nomaphila stricta (Acanthaceae), a newly c 

aquatic weed in Texas, and the first report for North America. Sida 15:11 5-117. 
Spencer, W. and G. Bowes. 1 985 . Limnophila and Hygrophila: a review and physiologic 

ment of their weed potential in Florida. J. Aquatic Pi. Managem. 23:7-16. 
Takematsu, T, M. Konnai, Y. Takeuchi and N. Ichizen. 1976. Weeds of cultivated f 

herbicides in China. Bull. Coll. Agric. Utsunomiya Univ. 9:91-107. 

AREA OF CENTRAL TEXAS— The Lacey oak, Quercus laceyi Small, is a small 
to moderate-sized tree that is commonly found on mesic north-facing slopes 
and in canyons on the Edwards Plateau of central Texas. The range of the species 
in Texas extends from Bexar and Medina counties in the east to Terrell and 
Brewster counties in the west; it is also found on the eastern slopes of the Sierra 
Madre Oriental in Coahuila and Nuevo Leon, Mexico (MuUer 1951, Nixon & 
Muller 1992). These plants were included in Q. glaucoides Mart. & Gal. by Tre- 
lease (1924) and Correll & Johnston (1970), but have recently been shown to 
represent a distinct allopatric species, with true Q. glaucoides being confined to 
central and southern Mexico (Nixon & Muller 1992). 

Quercus laceyt was described by Small (1901) from material collected "on lime- 
stone hills" in Kerr County, Texas, and subsequent descriptions of the habitat of 
the species (e.g., Muller 1951, Correll & Johnston 1970, Nixon & Muller 1992) 
have always emphasized the calcareous substrate. In October, 1993, 1 discovered 
a stand of several dozen individuals of g. lacey i in a mesic canyon at Enchanted 
Rock State Natural Area in Gillespie and Llano counties, Texas. The individuals 

Celtis reticulata Torr. and Diospyros texana Scheele. The following collection cita- 
tion is the first documentation of the occurrence of Q. lacey i on the igneous- 
derived sandy soils of the Llano Uplift rather than the limestone-derived soils of 
the surrounding Edwards Plateau: 

between Enchanted Rock and Little Rock, 1 3 Oct 1993, Lm/fee4l3S (SWT, TEX). 

Given that several vegetation studies have been conducted in the area now 
included within Enchanted Rock State Natural Area (Whitehouse 1933, 
Butterwick 1979, Walters 1980) and that the collection site is situated along a 
frequently used hiking trail, it is surprising that the occurrence of Q. laceyi has 

382 SiDA 16(2) 1994 

not been previously noted. This may be due to superficial similarities between 
Lacey oak and post oak (Q- stellata Wang.), which is common throughout the 
park. I myself have walked past this stand of trees numerous times over the last 
18 years and never noticed them until last fall. Upon close examination, how- 
ever, the characteristic bluish-green upper leaf surface and waxy deposits on the 
pulvinus readily distinguish Q. lacey i from Q. stellata. — David E. Lemke, Depart- 
ment of Bwlogy, Southwest Texas State University, San Marcos, TX 78666, U.S.A. 


BuriERwiCK, M. 1979. A surv. 

ey of the flora of Enchanted Rock and vicinity, L 

.lano and Gillespie 

counties, Texas. In Enchan 

ted Rock, a natural area survey (no. 14), pp. 4 

1-102. Lyndon B. 

Johnson School of Public/ 

iffairs, Univ. of Texas, Austin. 


^STON. 1970. Manual of the vascular plants oi 

f Texas. Texas Re- 

search Foundation, Renner 

, Texas. 

MuLLER,C.H. 1951. The oak; 

i of Texas. Contr. Texas Res. Found. 1:21-3 1 1 

Nixon, K.C. and C.H. Muller. 1992. The taxonomic resurrection ofQuera 

«/^.ej; Small (Fa- 

gaceae).Sida 15:57-69. 

Small, J.K. 1 90 1 . Trees and 

shrubs of the southern states-IV. Bull. Ton 

■ey Bot. Club 28: 


Trelease,W. 1924. The Ame 

Tican oaks. Mem. Natl. Acad. Sci. 20: 1-255. 

Walters, TW. 1980.Vasculi 

ir flora and vegetation of granite outcrops in tl 

le central mineral 

region of Texas. Masters thesis, Texas A&M Univ., College Station. 

Whitehouse,E. 1933. Plant; 

succession on central Texas granite. Ecology 1^ 


viarum Dunal, tropical soda apple, is a perennial shrub that belongs to section 
Acanthophora of subgenus Leptostemonum (Nee 1991). It is native to Brazil and 
Argentina but has become a weed in other areas of South America and in Africa, 
India, Nepal, West Indies, Honduras, and JMexico (Nee 1991) and Florida, U.S.A. 
(Coile 1993,Mullaheyetal. 1993a, 1993b, 1993c). 

Mature plants of 5. vtarum are 1 to 2 m tall and are armed on the leaves, 
stems, pedicles, petioles, and calyxes with broad based white to yellowish prick- 
les up to 12 mm long (Nee 1 99 1 , Mullahey et al. 1993c). Leaves and stems are 
pubescent; corollas are white with five recurved petals and white to cream col- 
ored stamens that surround the single pistil (Coile 1993). Immature fruits are 
mottled whitish to light green and dark green (i.e., like a watermelon) (iVIullahey 
et al. 1993c). The mature fruits are smooth, globular, yellow, and 2 to 3.2 cm in 
diameter with a leathery skin surrounding a thin-layered, pale-green, mucilagi- 
nous, scented pulp and moderately flattened, reddish-brown seeds (Coile 1993; 
Mullahey et al. 1993a). Each S. vtarum plant has the capability to produce over 
50,000 seeds (Mullahey et al. 1993c). 

The first known U.S.A. collection of 5'. viarum was from Glades County, Florida 

SiDA 16(2): 382, 1994 

Notes 383 

in 1988, but It may have been present in Florida as early as 198 1 or 1982 (Coile 
1993). Since its introduction into Florida, S. viarum has spread rapidly. By 1993 
it was reported to infest over 61 ,000 ha of improved pastures, citrus groves, 
sugar cane fields, ditches, natural areas (oak hammocks and cypress heads), etc. 
(Mullahey et al. 1993b). Drs. Robert Eplee and Randy Westbrooks (pers. comm., 
U.S. Dept. Agric, APHIS, PPQ, Whiteville, NC) estimated that by early 1994 
S. viarum infested as many as 285,000 ha in Florida. Because of its rapid popu- 
lation explosion in Florida, 5'. viarum was placed on the Florida noxious weed list 
on February 28, 1994. 

The primary means of dispersal of 5". viarum seems to be livestock and wild- 
life, such as raccoons, deer, feral hogs, and birds feeding on the fruits (Coile 
1993, Mullahey et al. 1993a). Mullahey et al. (1993a) reported that^. vtarum 
foliage is unpalatable to livestock although cattle will eat the mature fruit. Scari- 
fication of seeds by digestive systems of livestock and wildlife seems to promote 
seed germination. Intra- and inter- county and state movement of livestock that 
have recently fed on S. viarum fruit may be the primary vectors for its spread. 
However, contaminated hay may also serve as a means of dispersal (Wunderlin 
et al. 1993). Once S. viarum is established, livestock and wildlife continue to 
disperse seeds resulting in population levels that may increase exponentially. 

Collection data for S. viarum in Mississippi follow with herbarium abbrevia- 
tions after Holmgren et al. (1990), except MMNS and USMH (Mississippi 
Museum of Natural Science, Jackson and University of Southern Mississippi, 
Hattiesburg, respectively). 

Voucher specimens: U.S.A. MISSISSIPPI. Forrest Co.: NW of Hartiesburg, 5 mi NE jcc. of 
Hwys 1-59 & US 98, S of US 98, Southeast Mississippi Livestock Auction, 27 Jul 1994, Bryson 
13993 (BRIT/SMU, IBE, MISS, MMNS, NLU, PIHG, SWSL, USMH, VSC, VDB, additional 
specimens to be distributed); 1 5 Aug 1 994, Bryson 14188 & Rosso (SWSL, additional speci- 
mens to be distributed). Lamar Co.: S of Oloh, 2.8 mi S of Hwy US 98 and to E of Purvis-Oloh 
Rd, 27 Sep 1994, Bryson 14454, Byrd, & Rouse (SWSL, additional specimens to be distributed); 
Bryson 14456, Byrd, & Rouse (IBE, NY, SWSL, additional specimens to be distributed); Oak 
Grove Community, SE of jet. Oak Grove Road (= old Hwy US 1 1) & Griffith Rd, 27 Sep 1994, 
Bryson 1445 7, Byrd, & Rouse (SWSL, additional specimens to be distributed). Pearl River Co.: 
SE of Poplarviile, 4.3 mi SE jet. of Hwys 1-59 and MS 53, E ofMS 53, 14 Jul 1994, Byrds.n. 
(IBE, SWSL); 27 Jul 1994, Bryson 13996 (BRIT/SMU, DSC, IBE, MISS, MMNS, NLU, PIHG, 
SWSL, USMH, VSC, VDB, additional specimens to be distributed); 16 Aug 1994, Bryson 
14266 &Lockley (SWSL, additional specimens to be distributed); 8 Sep 1994, Bryson 14367 
(NLU, SWSL); SE of Poplarviile, 2. 1 mi SE jet. Hwys 1-59 & MS 53, N of Restertown Rd, 17 
Aug 1994, Bryson 14274 & ByrdiSWSL); SE of Poplarviile, 2.1 mi SE of jet. Hwys 1-59 & MS 
53, Eon Restertown, W of Bert Gentry Road, 17 Aug 1994, Bryson 14276 & Byrd (S^ii/SL). 
Stone Co.: ca. 1 1 mi WSW of Perkinston, 1.5 mi S of ,ct. Ridge Rd & Red Gap Rd on Red 
Gap Rd, 17 Aug 1994, Bryson 14279 & Byrd (SWSL, additional specimens to be distributed). 

The senior author first observed several plants of 5'. viarum growing in the 

median of Hwy I- 10 between mile markers 17 and 18 m Hancock County, 
Mississippi on 17 October 1993 but did not realize the significance of the obser- 
vation until the annual meeting of the Weed Science Society of America in 
February 1994. Subsequent trips to this site and surrounding areas in May, 
June, and early July 1994 were unproductive in relocating S. viarum because the 
median had been mowed. An additional sighting was reported from Hinds 
County, Mississippi in late 1993. 

At the Forrest County location six S. viarum plants were found along a live- 
stock holding -pen fence. Plants were about 1 m tall and were possibly estab- 
lished the previous year, each bearing up to 50 mature fruits per plant. About 
200 S. viarum plants in a 2 to 4 ha area were present at the inital Pearl River 
County location, ranging in size from seedlings to 1 .5 m tall with mature fruit. 
Additional seedling plants were observed with each successive visit to this loca- 
tion. At this site, plants were observed growing in the shade of pecan and oak 
trees but most were growing in full sun. Dead plants with mature fruit from the 
previous season were observed along a fence row that had been treated with a 
herbicide. The land owner stated that the plants had been at this Pearl River 
County location since 1991 and he pointed out several plants that had produced 
fruit in 1993. Evidently 5", viarum is at least a short-lived perennial in southern 
Mississippi. At the Stone County location, S. viarum infests several 100 ha of 
pastures and holding-pens around a farm headquarters barn at the end of Red 
Gap Road. In Lamar County, S. viarum infests a total of about 50 ha at two 
locations. At each of the locations in Lamar, Pearl River, and Stone counties, 
Solanum viarum was found in pastures where livestock had been introduced from 
Florida within the past few years and population size seems to be directly pro- 
portional to the total number of livestock imported from infested areas in Florida. 

At each location in Mississippi, plants were growing in a sandy loam soil. 
Mature plants were dug up at each location. In mature plants, S. viarum root 
systems were up to 3 dm deep with lateral roots up to 1 m long and 1.5 to 2.5 
cm in diameter. 

In laboratory and greenhouse experiments at Stoneville, MS, it was discov- 
ered that S. viarum fruit will ripen in sunlight if the green fruit were greater 
than 2 cm in diameter. Seed germination from these fruit was greater than 70% 
in greenhouse conditions. In addition to dispersal by livestock and wildlife, it 
was discovered that S. viarum fruit may be dispersed by water. Solanum viarum 
fruit from less than 1 cm in diameter to mature yellow or dry fruit are extremely 
buoyant. Thus, control oiS. viarum requires elimination of immature and ma- 
ture fruit as well as the whole plant including the root system. 

In order to prevent further dispersal of this pernicious weed, the U.S. Dept. 
Agric, APHIS, PPQ and the Mississippi Cooperative Extension Service are 

SiDA 16(2): 384. 1994 

initiating an education and notification campaign on the potential weed prob- 
lem of 5'. viarum. Additional surveys and an eradication program in Mississippi 
may be required to prevent S. viarum spread not only within Mississippi but 
into other states. Botanists, livestock producers, weed scientists, and regulatory 
officials should be alerted that S. viarum may already be in their region. Early 
detection is paramount to eliminate this weed, which has the potential to infest 
millions of ha of pasture lands, row crops, truck crops, sod farms, lawns, forests, 
and natural areas, especially in tropical and subtropical regions of the U.S.A. 
and other countries. Additional research is needed to determine S. viarmn fitness 
to more northern areas of the U.S.A. 

We thank Michael Nee (NY) for verification of the initial collections of 5". 
viarum and Richard Carter (VSC), Nancy Code (PIHG), Jeff MuUahey (SW 
Florida Research and Education Center, Immokalee, PL), and Robert Eplee and 
Randy Westbrooks (U.S. Dept. Agric, APHIS, PPQ, Whiteville, NC) for pro- 
viding information on the biology and spread of 5'. viarum in Florida. 
—Charles T. Bryson, USD A, ARS, Southern Weed Science Laboratory (SWSL), 
Stoneville, MS 58776, U.S.A. andJohnD. Byrd, Department ofPlant and Soil Sci- 
ences, Mississippi State Uniu, Mississippi State, MS 39762, U.S.A. 

CoiLE, N.C. 1993. Tropical soda apple, Solanum viarum Dunal: the plant from hell. Fla. Dept. 
Agnc. & Consumer Services, Div. Plant Industry. Bot. Circ. No. 27, 4 p. 

Holmgren, P.K., N.H. Holmgren, and L.C. Barnett (eds.). 1990. Index Herbariorum. Part 1: 
the herbaria of the world, 8th ed. New York Botanical Garden, Bronx. 

MuLLAHEY, J.J. and D.L. Colvin. 1993a. Tropical soda apple: a new noxious weed in Florida. 
Univ. Florida, Florida Cooperative Extension Service, Fact Sheet WRS-7, 3 p. 

MuLLAHEY, J.J., J.A. Cornell, and D.L. Colvin. 1993b. Tropical soda apple {Solanum viarum) 
control. Weed Technol. 7:723-727. 

MULLAHEY, J.J., M. Nee, R.P Wunderlin, and K.R. Delaney. 1993c. Tropical apple {Solanum 
viarum): ^v^eed threat in subtropical regions. Weed Technol. 7:783-786. 

Nee, M. 1991. Synopsis oi Solanum section Acanthophora: A revision of interest for glyco alka- 
loids, p. 258-266 in J.G. Hawkes, R.N. Lester, M. Nee, and N. Estrada, eds, Solanaceae III: 
Taxonomy, Chemistry, Evolution. Royal Botanic Gardens Kew, Richmond, Surrey, UK. 

Wunderlin, R.P, B.E Hansen, K.R. Delaney, M, Nee, and J.J. Mullahey. 1993. Solanum 
viarum and S. tampkense (Solanaceae): two weedy species new to Florida and the United States. 
Sida 15:605-611. 


Bremer, K. (with assistance of A. A. Anderberg, P. O. Karis, B. Nordenstam, 
J. Lundberg, and O. Ryding) 1994. Asteraceae: Cladistics and Clas- 
sification. (ISBN 0-88192-275-7, hbk). Timber Press, Portland, Or- 
egon. $79.95 (hbk). 752 pp., 89 figures, 24 tables, 6" x 9". 
Often viewed as a nontraditionalist for his early espousal of cladistic methodology, Kare 
Bremer has produced a monograph of the Asteraceae in the finest Benthamian tradition. This is 
no easy task in today's world of multiple sources of comparative data and conflicting taxonomic 
opinions among the disproportionately populous community of synantherologists. However, 
Bremer, who heads a Swedish research team in studying the systematics of the Old World 
composite tribes, has become a recognized authority on the Asteraceae as a whole. Bremer has 
created a handbook of genera and suprageneric taxa set in the context of insights from recent 
phylogenetic research and of unresolved problems. However, unlike recent symposium vol- 
umes which provide a forum for disparate views and often uneven treatments, this volume is 
the expression of relationships and classification seen from a single perspective. 

criteria, and 4) the origin of the family and major unresolved problems relevant to the family as 
a whole. Each of the systematic chapters provides 1) an overview of the variation in and past 
treatments of the subfamily or tribe, 2) a new or recently published cladistic analysis, 3) discus- 

Bremer's goal is "...not to provide a final classification of the Asteraceae, but rather to review 
the strengths and weaknesses of current classification to facilitate further revisions by future 
taxonomists" and "to provide a fairly even and consistent treatment of the tribes". In my opin- 
ion, he has succeeded in both. Even though the treatments of the Old World tribes are given 
somewhat more attention because the Stockholm group is most familiar with them, the dispar- 
ity does not create a noticeable unevenness, and the added insights are a plus for the North 
American synantherologist who has not ventured beyond the Isthmus of Tehuan tepee. 

In a sense, the value of the book depends on the audience. I believe that it will be of greatest 
use to students, to herbarium curators, and to amateur and general taxonomists. These people 
can use it as a reference to the genera of composites, as well as to understand the evolution of the 
family as whole. However, professional synantherologists, who already have references with 
generic descriptions, will find the value to be in the cladistic analyses and discussions. Bremer 
could have maintained the force in his views, as well as greatly reduced the size and price, by 
forgoing the generic descriptions. In fact, the information present is summarized in coded form 

Along this same line, one could criticize the cladistic analyses because the monophylesis of 

variable characters of certain large genera are coded for the most plesiom 
synapomorphic transition or whether the most plesiomorphic of a symplesiom 

if the segregate and the large potentially paraphyletic genus are treated in th 

However, the suprageneric classification is problematic in cases such as Dendroserh, which 
is treated in its own subtribe. Sanders et al. (1987) reviewed the suggestions for the origin 
of this oceanic island endemic. Included are Hieracium (probably sect. Stenotheca), Sonchus, 
Stephammeria, and a few other small genera. Bremer cites a paper that suggests the transfer 
of sect. Stenotheca to Crepis. Each of these four possible ancestors is in a separate subtribe. In 
Bremer's cladogram, Dendroseris is several branches and 10 steps away from Stephammeria, 
6 steps away from Crepis, and one step away each from Hieracium and Sonchus. Undoubt- 
edly, sect. Stenotheca should have been treated as a separate OTU and/or the large genera 
should have been represented by potentially monophyletic subgroups to more accurately 

Bremer and his associates have thoroughly referenced the pertinent literature. However, I 

,R.W.,T.F. Stuessy,C. 
n oiDendroseris and Robin 
\95-2l5.— Roger W. Sane 

and C. scoparia new to Mi 

Charles T.Bryson, John R.! 

Noteworthy new records for neotropical Gentiar 
Sagma ptocumbem {<Zi.r^op\vi\hctAQ), new to I 

iloptens thalictroides (Parkeriaceae) 


A new species of Zigadenus (Liliaccae) from New Mexico, ^ 
comments on tlie section Antklea 
William J. Hess and Robert C. Sivmski 

vith additii 

Una nueva especie ck 
Alvaro Cogollo P. ana 

;1 genero Pteropepon (Cucurbitaceae) 
'Johnj. Pipoly in 

de Colom 

John] Pipoly III 

xiccae: Caesaiipinioideae) from wester 

n Amazon] 

Anatomical study of Erioneuron and Dasyochloa (Poaceae: Chloride 

Eragrostideae) in North America 

Jesus Valdes-Reyna andStephan /,. Hatch 

on .he hybrid nature 




ONS TO Botany 

Lloyd H. Shinners 


Wm. F. Mahler 
Publisher 1971-1992 
Director Emeritus 

Barney L. Lipscomb 


Botanical Research Institute of Texas, Inc. 

509 Pecan Street 

Fort Worth, Texas 76 102-4060, USA 

817 332-4441 / 817 332-4112 FAX 

John W. Thieret Prof Dr Felix Llamas 

Associate Editor Contributing Spanish Editor 

Biological Sciences Dept Dpto de Botanica, Facultad de Biologia 
Northern Kentucky University Universidad de Leon 

lighland Heights, Kentucky 41076, USA E 24071 Leon, Spain 



Morfon Arboretum 

Lisle, IL 60332, U.S.A. 


Netu Mexico Forestry and Resources Conservation Division 

Santa Fe, NM 87304, U.S.A. 


'erstty of New Mexico 
,NM87131-1091, U.S.A. 

species chs 



ibed and illustratec 

estern New Mexicc 
the section Anticle. 

]. Its large flowers are 
mic, but quite locally 
). A discussion on re- 
? of the genus Z/^^i/f- 

describe e 

iluscra un; 



en,s nwgollomnsis. S 
ra manifiesto. Es er 

;us flores i 
'vo Mexic 

grandes son 
unque muy 
o. Se ofrece 

In 1968, Hess collected a Zigadenus from the interior of the Mogollon 
Mountains in southwestern New Mexico and identified it as Zigadenus 
virescens (Kunth) J.R Macbr. In the summer of 1993, he returned to the 
same general area and recollected this plant. Upon becoming familiar with 

misidentified and that this Zigadenus was undescribed. Through inquiry, 
Hess discovered that Sivinski was also studying this same plant in the 
Mogollon Mountains, and had independently arrived at the same conclu- 
sion that it was a new and undescribed species oi Zigadenus. 

Zigadenus mogollo 

nodis, 10-25(-35) x 3.5-8 c 


cepala elUp 

cica, margines saepe 

nrescentes (pallide fla' 

adaxialem > 

.ulgo pallidiores; api^ 

es) prope venam mediam abaxialem, 

, plana, breviter lanceolata, Capsulae crilobatae, 12-15 

Robust perennials, (4.5-)5.5-8.5(-9.5) dm tall. Bulbs slender, ovoid, 
2-3 X 1.3-2 cm, with thin, papery covering to thick, fibrous covering 
from old leaf bases, to 1 dm below ground. Basal leaves to 35 cm long, 10- 
18 mm wide, margins slightly scabrous; cauline leaves l-3(-5), reduced 
upwards. Inflorescences racemose, with (8-) 10-21 (-28) nodes, 10-25(- 
35) X 3-5-8 cm, occasionally paniculate with branches from lower 2 or 3 
nodes. Pedicels stout, 1.3-2 cm long, recurved at anthesis, erect in fruit; 
lowest bracts 2-4 x 0.5-1.2 cm, acute to acuminate, pale green, often 
purple-tinged along the margins, longer than pedicels, occasionally ex- 
tending beyond the flowers. Flowers campanulate and cernuous; tepals el- 
liptic to broadly elliptic, the margins often arching inward, 12-16 X 7- 
9.5 mm, purplish-red or margins purplish-red blending to pale yellow-green 
or green near midvein abaxial side, mostly lighter colored adaxial side; 
apices rounded, occasionally slightly emarginate; gradually tapering to base 
or broadly clawed; veins greenish, arching towards apex; glands 5-6.5 x 
3.2-5(-6) mm, slightly expanded terminally, apices emarginate or undu- 
late, somewhat bilobed, greenish-yellow; stamens with filaments 7-8 mm 
long, dilated at base, 1.5-2 mm wide, tapering to 0.5 mm wide at anther 
connection, anthers 2.4-2.6 x 1.7-2 mm, horseshoe-shaped, dehiscing 
entire length, pollen striate, monosulcate, 30-42p x 18-24}i; pistil 
tricarpellate, ovary partially inferior, trilocular, 3 styles and stigmata; ovules 
many, flat, short lanceolate. Capsules 3-lobed, 12-15 mm long, tepal lobes 
originating 4-5 mm above pedicels; no seeds seen. Flowering late August 
to early September. Fruiting September. 

392 SiDA 16(3) 1995 

Type: U.S.A. NEW MEXICO. Catron Co.: Mogollon Mcs., of the Gila Wilderness, Gila 

on Little Dry Creek Trail from Apache Springs, elev. 2860 m, 20 Aug 1 968, W. Hess 2212 
(holotype: MOR; isotypes: ARIZ; NCU; NMC; NY; OKLA; SMU; US). 

Additional collections examined: U.S.A. NEW MEXICO. Catron Co.: Mogollon Mts., 
S of Whitewater Baldy, center of sect. 30, Tl IS R18W, elev. 3l40 m, understory of dense, 
mature forest, 9 Sep 1980, R. Fletcher 4856 (UNM); 15 mi NE of Mogollon, 1 Aug 1938, 
C.L. Hitchcock, R.V. Rethke, R. van Raadshooven 4465 (WTU); along Bursum Road E of 
Mogollon, TllS R18W sect. 3, elev. 2770 m, 19 Jul 1994, C.A. Huff 1660 (UNM); 
between Apache Cabin & Sacaton, Baldy, Mogollon Mts., 9000 ft, 20 Jul 1959, A.R. 
Kruckeberg 4671 (WTU; UC); Gila Forest Trail 182, ca. 2.5 km S of NM 78, Tl IS R18W 
sections 2 and 11, elev. 3050 m, 2 Aug 1987, G.A. Levtn 1909 (NMC); Gila Wilderness, 

quent, elev. 3020 m, 2 Aug 1974, W.H. Moir 315 (NMC; RM); Mogollon Mrs., Forest 
Trail No. 182, Sandy Point, first 2 mi, ponderosa pine-aspen forest, 15 Aug 1974, D. 
Pinkava, E. Lehto & T. Reeves PI 2558 (ARIZ; WTU); Gila National Wilderness, Mogollon 
Mts., ca 10 air mi ESE of Mogollon and 1 mi from Hwy 78 on Trail 206 to Redstone Park, 
on W-slope, soil rocky clay with humus, Populus tumuloides Pteridiim and Rubus elev 
2680 m, 5 Sep 1976,7 Reitzel D Hill & R Spellenbeig 35 (NMC), Mogollon Mts , Forest 
Service Road 159 between Silver Creek Divide and Sandy Point, TllS R18W sect 3 
SW V.4, on rhyolinc soil, N-lacing slope in understory of mixed conifer forest of P'nudutsuga 

12 Aug'l993, R Sivinski & K Li^htfout 2317 (MOR, UNM), on NM Hw> 78 between 

Spellenberg & N Zmker 8237 (NMC), Mogollon Mts , Mogollon Road, 8 Aug 1900, E 
Wootons.n. (US). 

Zigadenus mogollonensis is apparently endemic to the mixed conifer and 
spruce-fir forests of the Mogollon Mountains between 2650 and 3200 meters 
in elevation. It is a common understory component of these high elevation 
forests and usually occurs on highly organic soils with a thick humus layer. 
Variation within this species is represented by its completely racemose or 
branched lower inflorescence, its range of tepal lengths between 12 and 16 
mm, and corolla pigmentation of pale yellowish-green with pale purple 
margins to green with dark brownish-purple margins. The exceptional speci- 
men o^ Reitzel et al. 53 (NMC) has green tepals with only little anthocyanic 
pigmentation, and is here placed within Z. mogollonensis because of its long 
(13 mm) tepals and campanulate corolla. Previous collectors have usually 
identified this species as Z. virescens, but also as Z. elegans Pursh and Z. 
porrifolius Greene. 

The nearest related species to Zigadenus mogollonensis are Z. elegans and Z. 
virescens. Both are widespread species, with Z. elegans ranging from Alaska 
to northern Mexico and Z. virescens occurring in the Sierra Madres of Mexico 
north to the southern mountains of Arizona and New Mexico. Zigadenus 
mogollonensis and Z. elegans have a similar inflorescence, which is usually 
racemose, but can have one or few lower panicled inflorescence branches. 

Rotate to rotate-campanulate corollas, erect pedicels at anthesis, and shorter 
(5—10 mm) tepals are characteristics of Z. elegans, whereas Z, mogollonensis 
is distinguished by its campanulate corollas, cernuous flowers at anthesis, 
and longer (12-16 mm) tepals. Zigadenm virescens is a more delicate plant 
with thinner pedicels and panicle branches, shorter bracts (usually less than 
the length of the pedicel), small (4-7 mm) tepals, campanulate corollas, 
and recurved pedicels at anthesis. Zigadenm mogollonensis has more robust 
pedicels and branches, larger bracts (as long or longer than the pedicels) 
and much larger tepals. The anterior margins of its tepals are also suffused 
with anthocyanic red over yellowish-green, which gives the flower a brown- 
ish-purple appearance. Both Z. elegans and Z. virescens frequently have 
anthocyanic bracts; however, this pigmentation infrequently extends to the 
flowers, and then is confined to the base of the tepals on the abaxial surface. 
Their open flowers are white, ochroleucous, or pale green. The references to 
purplish-flowered Z. elegans in western American floras (Correll & Johnston 
1970; Harrington 1964) are overstated and apparently derived from the 
description of purplish tepals for Z. coloradensis Rydberg (1900), which is a 
synonym of Z. elegans. The type specimen of Z. coloradensis is white-flowered, 
with small amounts of anthocyanic pigment on the abaxial surface of the 
tepals. The purplish pigmentation of Z. mogollonensis is much more intense 
and colors the anterior margins of the tepals. 

Zigadenus elegans is very rare in the MogoUon Mountains of New Mexico 
and is known from that range by a single 1881 collection {Rushy 406 US, 
only the flowering plant on sheet). However, Z, elegans is more common in 
the adjacent White Mountains of eastern Arizona and northern mountain 
ranges of Arizona and New Mexico. Zigadenus virescens is common in the 
MogoUon Mountains and occurs with Z. mogollonensis on the same forested 
slopes near Silver Creek Divide. No evidence of hybridization was observed 
at this point of sympatry. Pollen samples from four Z. mogollonensis and two 
adjacent Z. virescens were tested with Alexander's stain and found to be 99- 
100% viable. Late summer flowering times here are somewhat overlapped, 
but most Z. virescens flowered a little earlier and produced fruit at the time 
Z, mogollonensis was in full flower. A few unusual specimens of Z. virescens 
from the northern MogoUon Mountains {Daniel & Nelson 3598 ASU; Hubbard 
s.n. UNM) have large, leafy bracts that could possibly indicate a past cross- 
ing event with Z. mogollonensis. 

Zigadenus mogollonensis appears most similar to the large-flowered forms 
of Z, volcanicus Benth. in the Sierra de los Cuchumatanes in the Huehueten- 
ango district of western Guatemala. These two populations have long tepals, 
large bracts, cernuous/campanulate flowers and thick pedicels. They are 
differentiated by the somewhat larger, purple tepals of Z. mogollonensis com- 
pared to the white to ochroleucous tepals of the plants in Guatemala. 


The above species belong to the section Anttdea of the genus Zigadenus, 
which is characterized by a single bilobate or obcordate gland at the base of 
each tepal and a partially inferior ovary. Preece (1956) recognized five 
American taxa (Z. elegans var. elegans, Z. elegans var. glaucus (Nutt.) Preece, 
Z. virescem, Z. vaginatm (Rydb.) J.F. Macbr., Z volcamcus) in this section 
with one, or possibly two, additional species in Asia. Turner (1992) re- 
cently added another Mexican species (Z. hintoniorum B.L. Turner) to the 
section Antidea. The species of this section are usually distinct, but among 
the taxa intergradations and inconsistent variations are common. Species 
diagnosis must often rely on a combination of descriptive, qualitative fea- 
tures rather than definitive, measurable characteristics. Species descriptions 
that do not account for the range of variation have made this genus misun- 
derstood, and is evidenced by a long list of synonymy and the questionable 
rank of a few taxa. After studying the types and several hundred specimen 
sheets, the following discussion attempts to clarify some of the morpho- 
logical tendencies of the American taxa in the section Antidea oi Zigadenus. 

The two most important characteristics in separating taxa in the section 
Antidea are the position of the flower (erect or nodding) and flower shape 
(rotate or campanulate). The flowers at anthesis are either on erect pedicels 
with perianth widely spreading or rotate-campanulate, or cernuous pedicels 
with perianth consistently campanulate. Unfortunately, these characteris- 
tics are often difficult to assess on pressed and dried specimens, and collec- 
tors are well advised to note the floral aspects of fresh plants. It is also 
important to assess these characteristics on flowers in anthesis, since taxa 
with erect flowers can have cernuous buds, while those with nodding flowers 

Plants with erect/rotate flowers are best represented by the Ztgadenm 
elegans complex. This is the common North American species that extends 
from Alaska and eastern Canada to the southern Rocky Mountains and is 
sporadic in northern Mexico. It is very polymorphic, but retains some con- 
sistency by its erect pedicels and rotate corollas. Its flowers can often be 
attached to the erect pedicels at an angle that turns the faces of the flowers 
toward the outside of the inflorescence and perpendicular to its axis. This 
may give the false appearance of nodding flowers in some pressed speci- 
mens. Such minor and inconsistent variation is especially common in the 

from Colorado, Utah, northern Arizona, and Chihuahua. This variant of- 
ten, but not always, occurs in combination with very large bracts. 

The eastern variety, Zigadenus elegans var. glaucus, and a low elevation 
species from the canyonlands of western Utah, Z. vaginatus, are the only 

other erect pedicel-rotate corolla taxa presently recognized. The inflores- 
cence of Z. vaginatm is similar to Z. elegans, particularly with those variants 
from central New iVIexico that have a perpendicular flower attachment be- 
low the calyx. Most flowers of Z. vaginatm are often smaller, but within the 
range of typical Z. elegam. Cronquist et al. (1977) considered Z. vaginatm a 
synonym of Z. elegam. Preece (1956) and Welsh (1989) maintamed these 
taxa as distinct species because the former occurs at lower elevations in 
hanging garden seeps on canyon walls and has a later flowering period of 
July through September. Zigadenus elegam in Utah and Colorado grows at 
higher elevations and flowers earlier during June and July. Preece (1956) 
and Welsh (1989) stated that Z. vaginatm is closely related to Z. volcanicm 
in Guatemala. However, we believe it is more closely related to Z. elegam 
because of its erect pedicels and rotate corollas rather than to Z. volcanicm, 
which has cernuous/campanulate flowers. Zigadenm elegam also occupies 
similar habitats and tends to flower later south to the Guadalupe JVLoun- 
tains on the New Mexico/Texas border. A feature unique to Z. vaginatm is 
its numerous hard, persistent leaf bases, which allows it to be separated 
from Z. elegam. 

Zigadenm elegam is rare in northern Mexico, with a few known locations 
in Chihuahua and Coahuila. The Mexican populations differ from typical 
Z. elegam and have been treated variously. For instance, there is a large- 
flowered (tepals 10X5 mm long), large-bracted variant with rotate corol- 
las and perpendicular flower attachment on erect pedicels called Z. 
mohinoremis Greenm. Greenman (1903) does not mention flower angle, but 
it is evident on the type specimen {Nelson 4873 US). Preece (1956) placed 
Z, mohinoremis, known only from a few collections from Mt. Mohinora in 
Chihuahua, into synonymy with Z, elegam, but with some reservation. 

Zigadenm gracilentm Greene (1901) was described, based on plants col- 
lected in the Sierra Madre of Chihuahua {Pringle 1383 F, NY, US) and origi- 
nally identified as Z. elegam. They had campanulate corollas, tepals 6-7 
mm long, short stamens, and widely spreading pedicels. Greene described 

tics for the staminate and pistillate plants. There is some variation in tepal 
morphology; however, the types studied for this discussion had obvious 
stamens and maturing ovaries. Preece (1956) placed Z. gracilentm into syn- 
onymy with Z, elegam. In addition to the campanulate corollas and long, 
spreading pedicels, the flowers might have been cernuous at anthesis. The 
inflorescence bracts are narrow, acuminate, and 2/3 the length of the pedicels, 
and are identical to the long, Hnear-bracted form of Z, virescem that is com- 
mon in the northern Sierra Madres of Chihuahua and Sonora, and the 
Huachuca Mountains of southern Arizona. Zigadenm gracilentm is interme- 

396 SiDA 16(3) 1995 

diace between Z. virescem and Z. elegans and cannot be comfortably placed 
into synonymy with either species. The precise type locality of Z, gracilentus 
is unknown, but if it were to be relocated and the population had plants 
with consistently erect pedicels and campanulate corollas, it might be wor- 
thy of taxonomic recognition. 

South of the Rocky Mountains to Guatemala, the dominant form of Zi- 
gadenus has cernuous/campanulate corollas and is best illustrated by the 
variable Z. virescens. This species occurs predominantly in Mexico and usu- 
ally characterized by small tepals (4-7 X 1.5-3.5 mm), exserted stamens, 
short bracts, and slender, recurved pedicels. Zigadenus volcanicus, compared 
to Z. virescens, ranges further south into Guatemala and has slightly to greatly 
larger flowers (tepals 6-15 mm long), included stamens, and broader tepals 
(3-6 mm). Zigadenus mogollonensis is a New Mexico endemic with large 
purple flowers. 

There are many examples of herbarium specimens with inconsistent char- 
acter combinations for particular species. For instance, intermediate forms 
between Z. elegans and Z. virescens, within a fairly broad area of sympatry in 
eastern Arizona, southwestern New Mexico and northern Mexico, frequently 
have the tepal measurements, thicker pedicels and somewhat longer bracts 
of Z. elegans, and cernuous/campanulate corollas of Z, virescens. They are the 
dominant form in some localized areas (i.e., Mt. Baldy in the White Moun- 
tains of eastern Arizona). They also are in mixed populations since they 
share the same herbarium sheet with typical Z, virescens (i.e.. White Mts., 
A2, Peebles 12522 ARIZ) or with typical Z. elegans (i.e., Mogollon Mts., 
NM, Rusby 406 US). 

Several attempts have been made to recognize taxonomically these scat- 
tered, larger-flowered forms of Zigadenus virescens. The southwestern New 
Mexico plants were named Z. porrifolius Greene (1881), a name later ap- 
plied to several collections from eastern Arizona and an aberrant form from 
southern Coahuila (Hemsley 1885). In 1940, O. S. Walsh annotated many 
of these specimens as Z. virescens var. porrifolius, a combination that was 
never published. Kearney and Peebles (195 1) placed Z. porrifolius into syn- 
onymy with Z. virescens, but stated that it might be worthy of varietal 
status. Preece (1956) placed Z. porrifolius in synonymy with Z. virescens and 
attributed its larger flower size to the influence of Z, elegans. Zigadenus 
porrifolius is probably not worthy of taxonomic distinction because of its 
sporadic distribution and mixed populations. However, it is frequent enough 
to cause difficulties in species determination. 

In eastern and southern Mexico, Z. virescens may also intergrade with Z. 
volcanicus and possibly Z. elegans. There are several collections with long 
bracts, thick pedicels, and broader tepals of Z. volcanicus and Z. elegans, but 
with the distinctly decurved pedicels, campanulate corollas, and longer 

stamens of Z. virescens. A few larger-flowered collections from the border 
area between Coahuila and Nuevo Leon strongly resemble Z, volcankus, 
and the collection (McDonald 1322 TEX) from Coahuila could easily be 
placed within that species. This, and most of the tall forms of Z. virescens in 
eastern Mexico, have recently been named Z. htntontorum (Turner 1992). 
The holotype of Z. hintoniorum is somewhat aberrant, but it and the paratypes 
are clearly the highly variable species Z. virescens. In fact, the broad limits 
given to Z. hintoniorum would include Z. porrifolius and almost all of Z. 
volcanicus. Some Zigadenus from the Sierra Madre Orientale display odd 
character combinations that may be taxonomically divisible on a smaller 
scale; however, Z. hintoniorum is not correctly circumscribed and represents 
another attempt to segregate several intergrading and variable populations 
that cannot be comfortably assigned to Z. virescens., Z. elegans, or Z. volcanicus. 
Zigadenus volcanicus is a poorly understood species that has been incor- 
rectly represented by Preece (1956) and Turner (1992). Preece (1956) de- 
scribed Z, volcanicus as a short plant (2-3.5 dm) with erect or spreading 
pedicels and a corolla diameter of 1-1.4 cm, which could only be accurate 
if the corolla were rotate. However, among the specimens Preece annotated 
as Z. volcanicus, was one 7.5 dm tall {Seler 2323 US), and the type specimen 
{Hartweg 626 NY) was cut mto several sections to fit on the sheet! These, 
and all other specimens of Z. volcanicus seen in this study, were greater than 
3 dm tall and had cernuous/campanulate flowers. Turner (1992) inadequately 
characterized this species in his key to the section Anticlea and distinguished 
Z. volcanicus on the basis of tepals 14-15 mm long at anthesis. His descrip- 
tion was obtamed from a smgle Guatemalan specimen {Beaman 3097 TEX), 
which represented either an extreme variation or undescribed species with 
very large flowers that are not at all typical of Z. volcanicus. Duplicates of 
this unusual collection {Beaman 3097 US and MSC) have somewhat smaller 
tepals (12-13 mm long), less paniculate inflorescences, and a strong resem- 
blance to Z. mogollonensis. However, Z. mogollonensis is readily distinguished 
from it by its purple flowers and occurs 3,000 km north of the Guatemalan 

The type specimen and original description of Zigadenus volcanicus 
(Bentham 1842) represented a species with tepals 6-8 mm long. Standley 
and Steyermark (1952), from an independent collection {Steyermark 50133 
US) for their Flora of Guatemala, agreed with Bentham (1842). Baker (1879) 
recognized Z. volcanicus as a cernuous-flowered species and distinguished it 
from Anticlea mexicana Kunth (= Z. virescens) by its somewhat broader tepals. 
Many of the specimens of Z. volcanicus that we studied could be further 
distinguished from Z. virescens by their longer bracts (often as long or longer 
than the pedicels) and stamens that are shorter than the tepals. The com- 
bined features of broader tepals and longer bracts also describe Z. porrifolius 

and Z. hinuminriim (both = Z. virescens). Like Z. elegans, bract length was not 
consistent in Z. volcankus and varied from one-half to two times the length 
of the pedicel. Bract length in Z. virescens is also variable, but is usually less 
than the length of the pedicel. Other distinguishing, but inconsistent, fea- 
tures of Z. volcankus were noted in this study. Several specimens displayed 
a wavy, and often twisted, central axis of the inflorescence. The flowers 
were often clustered at the ends of long panicle branches. The significance 
of these characteristics must wait for the collection and study of additional 

The variation in Ztgadenus vokanicus suggests that it may be further di- 
visible, at least at an infraspecific level, and it is tempting to name the 
large-flowered form on the Sierra de los Cuchumatanes in northern Guate- 
mala. However, in several collections from that location, the tepals range 
in length from an extreme 15 mm to the 7 mm of the more typical form. 
One collection {Steyermark 30317 F) from Sierra de los Cuchumatanes is a 
mixed sheet containing a plant with tepals 10 mm long and another 7 mm 
long. Further collections and field studies are necessary before a large- 
flowered taxon can be separated from Z. vokankus. 

The following key to the taxa in Ztgadenus section Antklea accepts a 
broad concept of species variability and, therefore, somewhat artificial lim- 
its of taxon circumscription. The most ambiguous separation occurs be- 
tween Z. virescens and Z. vokanicus. Zigadenus hintoniorum will mostly key 
out to Z. virescens, but because there is no consistent and distinguishing 
criterion for Z. hintoniorum, it may also key to Z, vokanicus. We prefer to 
extend the limits of variation for Z. virescens and the geographic distribu- 
tion of Z. vokanicus rather than continue the use of a new and variable 
taxon that cannot be separated from existing species. Also, Z. porrifolius ( = 

We suspect it is the result of intergradation with Z. ekgans, and not be- 
cause of a relationship to Z. vokanicus. 


. Old leaf bases not persistent; tepals 5-10 mm long; plants wide.' 
3. Inflorescence usually racemose, occasionally with one or a fe 

1. Pedicels decurved or widely spreading, attached to cernuous flowers at an- 
thesis; corolla campanulate; plants of Arizona, New Mexico, Mexico, and 
4. Tepals 4-10 mm long at anthesis. 

5. Stamens usually as long or longer than the tepals; tepals 1.5-3.5(4) 
mm wide and < 8 mm long; floral bracts usually shorter than the 
subtended pedicels Z. Pirescem (Kunth) ] .7. Macbr. 

5. Stamens shorter than the tepals; tepals 3-6 mm wide and > 6 mm 
long; floral bracts V2 to longer than the length of the subtended pedicels 

Z. vokamcus Benth. 

4. Tepals 12-16 mm long at anthesis. 

6. Tepals greenish with pale to dark purple margins; plants of conifer 

forests in the MogoUon Mts. of SW New Mexico Z mogollonensis 


The senior author thanks Ian MacPhail for translating the description 
into Latin. Kim Altvatter and Nick Stoynoff accompanied him into the 
Gila Wilderness of the MogoUon Mountains for the field work, and he is 
eternally grateful. Floyd Swink did his superb editing for grammatical er- 
rors and Fayla Schwartz kindly reviewed the manuscript. The fine illustra- 
tion of this new species is by the hand of Nancy Barrels. The junior author 
thanks the curatorial staffs at ARIZ, ASC, ASU, CAS, DS, F, LL, MOR, 
MSC, NMC, NY, TEX, UNM, RM, US, and WTU for their prompt atten- 
tion to loans of specimens and type materials. He also appreciates the ef- 
forts of Guy Nesom at TEX, Steve Reed at UNM, and Susan Richardson at 
US in locating and providing several obscure pieces of literature. 

Baker, J.G. 1879. The aberrant tribes of Liliaceae. J 
Bentham, G. 1842. Plantae Hartwegianae. pg. 96. 


L, D.S. an 

d M.C. John 

STON. 1970. 

Manual of the vase 

ular pla 

nts of Texas. 


ai-ch Foun 

er, Texas. 



N, N.H. Hoi 
e 6. The Ne- 

,MGREN,J.L. Reveal: 

and PK 
irden, E 




U. New spec 

les of plants. 

, chiefly New Mexic 

an. Bui: 

I. Torre> 





)teworthy sp 

ecies XXVIII. Pitto 



.AN, J.M. 

1903. New 

and otherwi 

se noteworthy angi 


; from I 



ca. Proc. Am, 

.r. Acad. Arts 39:69-120. 

jGTON,H.D. 1964. Mar 


ants of Colorado. S'^ 

callow Press, Inc 



Y, W.B, 1 


L Centrali-Ar 

nericanae. Botany 3 


lY, T.H. a 

tnd R.H. Peebles. 1951. 

Arizona flora. Univ 

'ersity c 

)f Califc 



toral Dissertation, Washington State University, Pullman. 
Rydberg, p. a. 1900. Studies on the Rocky Mountain flora— II. Bull. Torrey Bot. Clu 

Standley, PC. and J. A. Steyermark. 1952. Flora of Guatemala. Field Mus. Nat. Hist 

Bot. ser. 24(3). 

Welsh, S.L. 1989. On the distribution of Utah's hanging gardens. Great Basin Naturalis 


De Waal, Louise C, Lois E. Child, P. Max Wade, and John H. Br 
(Eds.). 1994. Ecology and Management of Invasive Riverj 
Plants. (ISBN 0-471-94257-X, hbk.) John Wiley and Sons L 
Chichester, West Sussex, P019 lUD, England. $95.00. 217 pp. 

ent habitats are invaded by non-indigenous plants, I was somewhat intrigued wh) 
riverside habitat was chosen. The preface provides scant information on this problem 
only clue offered is that the river environment is "especially prone to invasion." An 



Fundactonjardm Botdmco, "Joaquin Antonio Vribe" 

Apartado Aereo 31407 

Medellin (Antioquia) COLOMBIA 


Botanical Research Institute of Texas 

309 Pecan Street 

Fort Worth, TX 76102-4060, U.S. A. 

Pteropepon oleiferum i 


bed, illustrated, and Its 





'ithin tl 

he genus and family, bas 


uher on a conic fila 

ment, c 

:oncomitant with its d 

loecious habi 

t. Nev 


;, we feel 

lat the new entity i; 

s best I 

placed in Pteropepon than in any otht 

;r cuct 


us genus 

nown, because of Its J 


^compressed, fibrous an 

.d winged frui 

it, and 

staminate flowers 

1 panicles. As a resul 

t of the 

• authors' collaborative 

research, this 

IS the 

third ne- 

w species 

) be described from 

the Rfc 

iClaroareaof Antioqu 

ia since 1991 

. Thei 

irea is ca 


biogeographically interesting. 

El genero Pteropepon (Cogniaux) Cogniaux (1916) fue descrito para incluir 
las especies de Skydium Schlechtendal que son monoicas, con Acres 

o subsesiles; frutos lateralmente comprimidos, fibrosos y alados. Las otras 
especies que quedaron en Skydium son dioicas, con flores estaminadas y 
pistiladas en paniculas, el androceo de 2 anceras con 2 tecas y una con una 
teca, filamencos pequefios, y bayas globosas. En Das Pflanzenreich , Cogniaux 
reconocio dos especies, ambas del sureste de Brasil. Veinte afios despues, 

SiDA 16(5): 401 -406. 1995 

402 SiDA 16(3) 1995 

Macbride (1937) describio una variedad de Pteropepon deltoideus var. peruvianm, 
segun las determinaciones de Harms, para enronces llegar a un total de 3 
taxa en ei genero. Desde entonces, Jeffrey (1978) y Kearns (1993) no han 
reconocido yd.t. penwianm como entidad meritoria, aunque no han explicado 
el porque. Adicionalmente, Crovetto (1950, 1952) describio Pteropepon 
parodit and P. argentinkme, respectivamente, y creo la seccion Mkropteropepon, 
para incluir las especies que tienen frutos pequenos y membranaceos, 
incrementando el total de los taxa hasta cinco. Como resultado de las 
investigaciones en la region de Rfo Claro en el Valle del Rio Magdalena en 
Colombia, encontramos una especie que se ubique mejor en este genero 
que en cualquier otro. La especie es otro ejemplo del alto nivel de endemismo 
en esa zona, como fue el caso con otras especies pertenicientes a los generos 
Erythroxylum (Cogollo & Pipoly 1993) y Cybianthus (Pipoly 1991). La nueva 
especie aqui descrita se separa de las especies previamente conocidas por la 

^ Lamina membianacc.i, ci lan^'ulai o subdclroidca, apict acuminado, base 

papiiosos; frutos costados P. mvnospernws (.Veiiosoj Cogniaux 

4. Sepalos subulados; petalos anchamente ovados, glabros; frutos no 

costados P. deltoideus (Cogniaux) Cogniaux 

3. Lamina coriacea, ovada o oblongo-eliptica, apice agudo, base 

redondeada; estambre l;anteras biloculares P oleiferum Cogollo and Pipoly 

Pteropepon oleiferum Cogollo and Pipoly, sp. nov. (Fig. 1) 

Trepadora dioica; tallos graciles, surcados, glabrescentes, ramificado: 

coriaceas, ovadas a oblongo-elipticas, (6.5-)7.5-l6.0(-17.0) cm de largo, 
(4.5-)8.5-10.0(-13.2) cm de ancho, el apice agudo, la base redondeada, 
glabras por ambas caras, las nervaduras laterales 3 a cada lado, impresas a 

rior, el margen entero; zarcillos graciles, redondos a ligeramente surcados, 

404 Si DA 16(3) 1995 

bifidos en el apice. Inflorescencias escaminaclas paniculadas; pedunculo prin- 
cipal plurifloro, gracil, surcado, glabro a ligeramente papiloso, 3.5-30.0 
cm de largo, las ramas divergences, las basales de 2.5-12.0 cm de largo, 
mas cortas hacia el apice, subtendidas por unas bracteas de 2.0-7.0 mm de 
largo. Flores escaminadas pequeiias, verdosas, 2.0-2.5 mm de diametro; 
receptaculo glabro, rotado; sepalos 5, oblongos, atenuados en el apice, 
uninervios, abaxialmente papilosos, adaxialmente glabros; corola dividida 
hasta el receptaculo; petalos 5, ovados, glabros, alternandose con los sepalos, 
un poco mas grandes que los sepalos; escambre 1, pequei^o, el filamento 
conico, de 0.3-0.5 mm de largo, la ancera 1, amarilla, bilocular, loculos ca. 
0.5 mm de largo, 0.3 mm de diametro. Flores pistiladas desconocidas. Fruto 
fibroso, coriaceo, eliptico a obovado, glabro, 9.0-10.5 cm de largo, 6.0-8.0 
cm de ancho y 2.0-2.5 cm de espesor, alado, con 2 costillas prominentes 
longitudinales. Semilla 1, eliptica, de 5.0-6.0 cm de largo, 4.0-5.0 cm de 
ancho, verrugosa, con testa lenosa. 

Tipo COLOMBIA Ancioquia Municipio de Puerto Triiinfo, autopisc.i McJellin-S.incafe 
de Bogota, sector Rio Claro-Rio Magd.dtna (.imino de l.i autopisca hacia la griica de "El 
Condor," ca 400 m de la autopista ' 0^ V, \i i ' Mi'W o()-400 m, 27 Feb 1994 (fl 
estam), /i Coy^ollo & G. Jcjrcjmillo Sn.'Ji |^\n\un\^^ )Al M im)iik)s BRIT, COL, IIUA, 

Paracipos COLOMBIA Atitioquia Muiuupiodc \.n I iiis, ( anon de Rio Claro, margen 
dereclia, sector sur, 05' 5 ^ N, ^ I" ^'/W ->2=i- 1 1 m, K)c t I 98 ■, (fr), A (j>g<,U<, & R Borju 
764 (COL, JAUM. MO), ^ Die 1 9S ^ (ll csram ), .\ Coi^'J/o 1040 (COL. JAUM, MO), 

Dtstnbiaion. — Pteropepon oleiferum es conocida hasta ahora unicamen 
de la region de rio Claro, en las faldas onent.iles de la Cordillera Central c 
los Andes, hacia el valle medio del Rfo Magdalena, en los municipios c 
San Luis y Puerto Triunfo, Departamento de Antioquia, Colombia, ent 
300 y 700 m de altitud. 

Etimologia. — El epfteto especifico "oleiferum" es del latfn, "oleum" (aceit 
con el sufijo adjectival "-fer" (de portar, o contener); asi que se refiere 
hecho de que la semilla contiene aceite. 

Ecologia. — P. oleiferum crece en orillas de quebradas, en bordes de boscju 

por agua. 

Usos y Condiciones Actuales en Cuanto a Conservacion: Las semillas c 



PiPOLY, A new species of Pteropepon from CoL 





. una gran cantidad de aceit 

e, el cual es 

extraido en 


1 rudii 

Tientaria ] 

3or algunos habit 

antes de la 

region, y ej 

i usado para 



de herran 

aientas. La genre: 
los lugares hay pi; 

local apreci 
anes de dor 

a el valor de 

la especie y 


wopepon oleiferum 

es una especie u 

nica denrrc 

) del genero 

, por su fior 



con un sol 

esrambre, anrera 


nsertado en i 

an filamento 


:o, y por su co 

ndicion dioica. S 

;m embar^ 

;o, debido i 

1 sus frutos 



te comp 

rimidos, fibrose: 

5 y alados 

■ flores est: 

aminadas y 



,, P. oleifer, 


comodada i 

.n otro gene 

ro conocido. 



e de la z, 

ona del Rio Clare 

) sobre roc: 

contiene no 

solamente las tres nuevas especies susodichas, sino otra medio docena de 
entitades no identificables hasta la fecha y posiblemente nuevas tambien. 
Estudios actualmente progresando indica que la zona tiene una mezcla rara 
de elementos chocoanos, andinos, amazonicos y centroamericanos, 
verificando su importancia como fuente de biodiversidad unica en el 
departamento de Antioquia. 


La colaboracon de nuestros compaiieros del herbario respectivos se ha 
facilitado el trabajo para ambos autores, y les agradecemos. En particular, 
el trabajo de Dayron Cardenas (CO A), Luz Marina Velez, y Juan Guillermo 
Ramfrez nos facilito mucho en compilar los datos sobre la especie. Lindsay 
Woodruff revise el manuscrito y hizo comentarios valiosos. Apreciamos el 

Investigaciones Cientificas y Proyectos Especiales, "Francisco Jose de 
Caldas"-COLCIENCIAS lo que ha permirido realizar exploractions en la 
region de Rio Claro; tambien el apoyo de la National Geographic Society, 
cuya subvencion para estudios en el Parque Nacional Natural "Las 
Orquideas" nos ha permitido reunirnos bianualmente tanto para estudios 
de la flora del parque como estudios sobre regiones adyacentes, como el 
presente. Tambien, comentarios sobre el manuscrito, hechos por Michael 
Nee (NY), Charles Jeffrey (K), y Dennis Kearns (MO) ennquecieron y 
mejoraron la presentacion. Apreciamos el trabajo de la dibujante Consuelo 
Garcia (HUA) por la ilustracion aqui presentada. 

EFFREY, C. 1978. Fun 

:earns,D. 1993. Cue 
logue of the floweri 

has subgenera Conomorpha (Myrsinaceae) 


ki, Dorothy VanD^kl John Bi nimmin Li aki , and Marc i lotil Li a 
RoLDLR 199? Desert and Mountain Plants of the South we; 
{ISBN 0-8{)6l-2489-x, pbk) University of Oklahoma Press, PO 

787, Norman, OK ^^070-0787 $18 95 2^9 pp 





Botanical Research Institute of Texas 

309 Pecan Street 

Fort Worth, TX 76102-4060, U.S.A. 

all, T. vasquezit. 

zits oiTachtgali,T i 


Tachigali Aublet is a neotropical genus now considered to contain ap- 
proximately 60 species, including the 35 species formerly included in 
Sclerolohium Vogel (Zarucchi & Herendeen 1993). Historically, the genera 
were distinguished by position of the pistil stipe with reference to the re- 
ceptacle cup iSclerolobium, centrally; Tachigali, eccentric) more than any other 
character (Dwyer 1954, 1957). However, Zarucchi and Herendeen indi- 
cate that there is a continuum m the stipe character and no other either 
morphologcial or anatomical character may successfully be used to separate 
the two groups. Both are frequently myrmecophilous, a trait otherwise rare 
in the Caesalpinioideae. 

While carrying out a series of florulas in the western Amazon Basin of 
Peru and Colombia, a new taxon was discovered which is assigned to 
Tachigali and described herewith. 

Tachigali vasquezii Pipoly, sp. 

praeclare distat. 

Emergent canopy tree to 30M5) m tall, 57 cm DBH; branchlets subterete 
to 3-5-anglecl, bark brown and gray in alternating longitudinal bands, 
0.8-1 .2 cm diam., minutely chocolati-tomentellous, early glabrescent; pith 
large, soft; stipules foliaceous, bifoliolate, coriaceous, ovate, 1.8-3.5 cm 
long, 1.3-2.8 cm wide, apex acute, base asymmetric, truncate, midrib im- 
pressed above, prominently raised below, secondary veins 5-9 pairs, buUate 
and densely chocolati-tomentose and glandular along the midrib and sec- 
ondary veins above, densely chocolati-tomentose below, the margin entire, 
revolute, glabrous. Leaves paripinnate, alternate, petiolate; petioles (5-)9- 
l4(-27) cm long, 0.7-1.5 cm diam., pulvinate basally; rachis solid, 
subterete, with an adaxial furrow, without myrmccodomatia, densely and 
minutely chocolati-tomentellous, (l4-)24-36(-47) cm long, apically mu- 
cronate, the mucron 1-2 mm long, caducous, eglandular; petiolules terete, 
(0.7-)0.9-l(-1.3)cm long, 2-3 mm diam., densely chocolati-tomentellous, 
the tomentum persistent; leaflets coriaceous, oblong, 12-18(-31)cm long, 
5-7 (-13) cm wide, apex abruptly acuminate, the acumen 0.5-1 cm long, 
base truncate, midrib impressed above, prominently raised below, densely 
tomentulose, secondary veins l4-20(-28) pairs, densely tomentellous and 
somewhat impressed above, prominently raised and tomentose below, sub- 
marginally loop-connected to the next distal secondary vein, tertiary veins 
somewhat impressed above, prominently raised below, nitid above, densely 
chocolati-tomentellous-velutinous below, the margin revolute, entire. In- 
florescence an axillary panicle, 26-29 cm long, peduncle 6-9 cm long; 
pedicels 3-7 mm long. Flowers unknown. Fruit flat, oblong or rarely ellip- 
tic, 12.5-15 cm long, (3-)5-6.5 cm wide, apex rounded, base acute, testa 
reddish-coffeate, exfoliate at maturity, one-seeded. 

Type. PERU. Amazonas: Valle del Rio Santiago, 65 km N of Pinglo, Quebrada Caterpiza, 
2-3 km behind Caterpiza, 200 m, 1 Feb 1980 (fr), V Huashtkat 1910 (holotype: MO; 

Amacayacu, Quebrada Agua Pudre, 1.5 km NE of Rio Amacayacu mouth, MO Strategy 
Inventory site, tree No. 158, 03°47'S, 70°15'W, 200-220 m, 11 Nov 1991 (ster.),/ 
Pipo/y eta/. 15851 (BRIT, COL, FMB, MO). PERU. Huanuco: Prov Pachitea; Codo de 
Pozuzo; alluvial floodplain of Rio Pozuzo after it emerges from mountains, trail N of 
settlement to Rio Mashoca, 75° 25'W, 9° 37'S, 500 m, 19 Oct 1982 (ster.), R. Foster 9329 
(F, USM). Loreto: Prov. Maynas, Distrito Iquitos, Allpahuayo, Estacion Experimental del 
Instituto de Investigaciones de la Amazonia Peruana, 04° 1 0'S, 73°30'W, 1 50-1 80 m, Dec 

1990 (ster.), R. Vdsquez & N.Jaramillo 13634 (AMAZ, MO, USM). 
Manu, Parque Nacional Manu, Rio Manu, Cocha Cashu Station, 350 m, 23 Nov 1980 
(ster.), R. Foster 5830 (F, USM), Pakitza Station, Rio Manu, North Trail, 350 m, 20 Nov 
1980 (fr), R. Foster 5780 (F, USM), Zone 1, one km N of camp, 11° 56'S, 71° 16'W, 350 
m, 22 Dec 1988 (ster.), R. Foster & S. Balckon 12671 (F, USM). Prov. Tambopata, Tambopata, 
along Ri'o Tambopata, 12°49'S, 89°18'W, 280 m, 19 Feb 1984 (ster.), A. Gentry et al. 
45633 (AMAZ, CUZ, MO, USM); Tambopata Tourist Camp, at jet. of Rios Tambopata 
and La Torre, 12°49'S, 69°43'W, 280 m, 22 July 1985 (ster.), A. Gentry et al. 31088 
(CUZ, MO, USM), 26 May 1987 (ster.) A. Gentry et al. 37661, 30 May 1987 (ster.), A. 
Gentry et al. 37948; Las Piedras, Cusco Amazonico, Permanent Inventory Plot, 12°29'S, 
69°03'W, 200 m, 25 Nov 1991 (ster.), M. Timand &N.Jaramtllo 3464 (CUZ, MO, MOL, 
USM), 31 Oct 1991 (ster.), M. Timand & N. J aramillo 2934 (CUZ, MO, MOL, USM); 18 
Jun 1989 (ster.), 0. Phillips etal. 412 (CUZ, MO), 2 1 Jun 1989 (ster.), 0. Rhilltps et ai 553 
(CUZ, MO, USM). Pasco: Prov. Oxapampa, Valle del Palcazu, Iscozacin, camino a Villa 
America, 400 m, 8 Aug 1981 (ster.), R. Foster 4688 (F, USM); PEPP Arboretum, 09° 50'- 
10°45'S, 68° 00'-68° 30'W, 300-600 m, (star.), G. Hartshorn, J. Qiujano & E. Meza 2871 
(CR, F, US, USM); Palcazu, Rfo Alto Iscozacin, Ozuz to Rio Lobo, 10° 19'S, 75°l6'W, 10 
May 1985 (fr), R. Foster & B. d'Achille 10070 (F, USM). 

Common ;?^wt,— "chaira pacae amarlUa" (Peru). 

Dtstribution: Throughout the western Amazon Basin, from near Leticia, 
Colombia, south and westward through the departments of Loreto, 
Amazonas and Pasco, to Madre de Dios, Peru, at 150-600 m elevation. In 
addition, reported to be common in the departments of Pando, Northern 
La Paz, and Beni, Bolivia (R. Foster- pers. comm.). 

Ecology. — Common in the canopies of humid forests, on mostly old allu- 
vial clays, especially sandy clays, or rarely, sands. T. vasquezii is frequently 
encountered on non-inundated terraces of old floodplains and low hilltops 
and slopes of the Upper Amazon and Andean foothills. Quantitative forest 
inventories containing permanent plots, established near Leticia, Iquitos 
and Cusco Amazonico have shown that 3-5 individuals of T vasquezii may 
be found per hectare, sometimes clumped. Other data from those plots 
have revealed that T. vasquezii regularly grows in the same habitat as T. 
ptychophysca Spruce ex Bentham, but the two do not grow in close proxim- 
ity. According to Robin Foster (pers. comm.), this species is monocarpic- — 
individuals flowering once and then dying slowly as the seeds mature. The 
behavior is similar to that described for T. versicolor (Foster 1977), except 
there seems to be greater frequency of individuals in which only a part of 
the tree flowers and dies within a given year. 

Etymology. — This species is dedicated to Rodolfo Vasquez Martinez, 
friend, colleague and director of the Peruvian exploration project for the 
Missouri Botanical Garden. He is principal author of the Florula of the Bio- 
logical Reserves of Iquitos, containing nearly 3,000 species. He is also an au- 
thority on the systematics of the Hypericaceae, Ebenaceae and Myrsiticaceae 
of Peru, and the genus Caraipa (Clusiaceae) throughout its range. 

The angulate branchlets, coriaceous leaves with acuminate apices and 

stipules, indicate that Tachigali vasquezii is most closely related to the 
vicariant T. rugosa (Martius) Pipoly and Zarucchi of the cerrado formations 
from eastern Brazil. However, T vasquezii is easily recognized by the choco- 
late tomentum of the branchlets and abaxial leaflet surface, smooth oblong 
leaflets, and terete, longer petioles. Whether it is coincidence that the re- 
cently described Affonsea Pipoly and Vasquez, is also most closely related to 


As was previously mentioned in the introduction, Tachigali and 
Sclerolohium cannot be separated (Zarucchi & Herendeen 1993) on any char- 
acter other than relative position of pistil stipe within the receptacle cup. 
The plasticity of this character, and lack of any other known character to 
separate the two groups necessitates the transfer of three species of 

PoPOLY, Tachigali I 

Tachigali rugosa (Martius) 

Sclerolobtum rugomm Martius t 

BRAZIL. MatoGrosso: Cni^h^,daSilva Mamo s.n., Martius herbarium 1155. 

lotype: M (not seen); photo at F Neg. 6264). 
Tachigali micropetala (Ducke) Zarucchi and Pipoly, comb. nov. Basic 

Sclerolobtum micropetalum Ducke, Bol. Tech., Inst. Agron. Norte Belem. Type. I 

ZIL. Amazonas: Manaus, without date, Ducke 1219 (syntypes: K, MO, NY). 
Tachigali bracteosa (Harms) Zarucchi and Pipoly, comb. nov. Basic 

Sclerolobium bracteosum Harms, Verhandi. Bot. Ver. Brandenb. 48:168. 1907. ' 
leiras, Ule 6094 (F- fragment, I 

ducted during my tenure at the Missouri Botanical Garden, under the aus- 
pices of the John D. and Catherine T. MacArthur Foundation. Supplemen- 
tal research on the Flora of Peru, which also contributed to fieldwork, was 
carried out through a grant from the Andrew W. Mellon Foundation. 
Rodolfo Vasquez, who provided the excellent illustration, is the director of 
the Peru program at the Missouri Botanical Garden, and is also supported 
by grants from the John D. and Catherine T. MacArthur Foundation for 
permanent inventory plot studies and the Andrew W. Mellon Foundation 
for the Peruvian exploration program. I thank Jim Zarucchi (MO), Alex 
Lasseigne and Robin Foster (F) for their reviews and helpful suggestions 
concerning the manuscript. 

1954. Rapports entre stipe et coupe receptaculare dans la classification des 

lerstieae (Caesalpmiaceae). VIII. Cong. Int. Bot. Rapp. and Comm. Sect. 4:52-54. 

1957. The tropical American genus Sclerolobtum Vogel (Caesalpiniaceae). 


7. Tachtgat 

ta versicolor is 

a suicidal 

neotropical tree. Nature 268(5621 

J. a. 

id P. Hhrei 

.), Catalogue 
;ouri Bot. Gai 


(Fabaceae). Pp. 1254-1255. In: L. 
wering plants and Gymnosperms o 


Powell, A. Michael. 1994. Grasses of the Trans-Pecos and Adjacent 
Areas. (ISBN 0-292-76553-3, hbk; ISBN 0-292-76556-8, pbk). 
University of Texas Press, PO. Box 7819, Austin, TX 78713-7819. 
$75.00 hbk; $29.95 pbk. 377 pp. 

peaks a mile or more high (up to 875 1 ft on Guadalupe Peak). Powell's book accounts for 
83 genera and 268 species of Trans-Pecos Poaceae (half of the grasses known from Texas). 
Endemic to that part of the state are 53 species. Most of the illustrations are the by-now- 

i chapters te. 
>ans-Pecos I 

;ra and species are at appropria 

ous taxa. Nomenclature is up to date.^ohn W. Thieret. 

Jones, David L. 1993. Cycads of the World. Ancient Plants in Today's 
Landscape. (ISBN 1-56098-220-9, hbk). Smithsonian Institution 
Press, 470 L'Enfant Plaza, Washington, DC 20560. $45.00 (plus $2.25 
postage). 312 pp. 

This is a one-volume encyclopedia devoted to the 185 species of cycads. The illustra- 
tions — of plants, leaves, and cones — are the book's principal glory. Many (ca. 250) are fine 
color photos, some are original line drawings, and a few are reproductions of old engrav- 

—e.g., gametophyte development, anator 

1 genus and species. Generic accounts inc 
given for Cycas, Encephalartos , and Zamia, 

nd propagation. Species accounts includi 






Departamento de Botdnka 
Universidad Autonoma Agraria "Antonio Narro" 
Buenavtsta, Saltillo, Coahmla 25513, MEXICO 


S.M. Tracy Herbarium 

Department of Rangeland Ecology and Management 

Texas A&M University 

College Station, TX 77843-2126, U.S.A. 

.eaf blades of Eriomuron and Dasyochloa species were analyzed for anatomical de 

I. Differences between E. avenaceum and E. grandiflorum were not found, supporting 
;rpretation that these are conspecific. Differences between E. avenaceum, E.pilosum 
'ealleyi were found, suggesting that these be maintained as distinct entities. Differs 
ween Dasyochloa and Erioneuron support the recognition oi Dasyochloa as a monoi 
rth American genus. 



as la 
do. I 

[minas de 
li analisis 

las especies de Ertonei 
de la lamma en secc 


y Dasyochloa, par 

a epider 







. No se encontrarc 

in diferei 




um, lo que apoya la int 







Caceres (1950) studied the leaf blade anatomy oi Munroa mendocina and 
Blepharidachne benthamiana Hitchc. and compared these species with spe- 
cies oiEragrostis and Tridens pilosa (Buckl.) Hitchc. var. argentina (O. Kuntze) 
L.R. Parodi [syn. Erioneuron pilosum (Buckl.) Nash var. longiaristatum (Kurtz) 
Anton}. Tateoka (1961) examined the leaf epidermis oi Tridens, including 

Chloridoid subtype as described by Prat (1936). 

Sanchez (1979a) studied the leaf blade anatomy of three species ofTridens 
and two species, including their varieties, of Erioneuron from Argentina. 
Observations of the unique anatomical features of Dasyochloa pulchella 
(H.B.K.) Willd. ex Rydberg [syn. Erioneuron pulchellum (H.B.K.) Tateoka} 
were made. Sanchez (1979b, 1981) reported the development of the "Kranz" 
structure in the stems oi Munroa, Blephandachne, and Erioneuron from Ar- 
gentina. Sanchez (1983) analyzed the anatomical differences between 
Dasyochloa and Erioneuron and proposed the segregation oi Dasyochloa as an 
independent genus following, in part, criteria as outlined by Parodi (1934). 

The grass genus Erioneuron was described by Nash in Small's (1903) 
Flora of the Southeastern United States, based on Erioneuron pilosum. Tateoka 
(1961), in a biosystematic study of the genus Tridens, presented cytologi- 
cal, morphological, and anatomical differences to support recognition of 
two distinct genera. One prominent difference was in the basic chromo- 
some numbers, x == 8 for Erioneuron and x = 10 for Tridens. The results of 
the examination of the leaf anatomy show that all species of Tridens and 
Erioneuron have a Chloridoid subtype of the panicoid type of leaf epidermis, 
characterized by the presence of globose or club-shaped bicellular microhairs 
and saddle-shaped silica bodies. Particular kinds of bicellular microhairs 
were observed in Tridens and Erioneuron. Tridens had bullet-shaped hairs, 
and Erioneuron had ellipsoidal hairs. Differences in the leaf cross-section 
were found between genera in leaf margin, midrib of the leaf blade, and in 
the number of vascular bundles within the leaf. Based on these results 
Tateoka recognized the genus Erioneuron and suggested affinities with the 
genus Munroa, rather than with Tridens. Sanchez (1979b) studied the leaf 
anatomy of the species and varieties of Tridens and Erioneuron from Argen- 
tina. That study included a table of characters used to separate the genera, 
with keys to the species and varieties based on anatomical characters. This 
classification has been well accepted by agrostologists and used in several 
grass floras (Correll & Johnston 1970; Nicora 1973; Gould 1968, 1975, 
1979; Anton 1977; and McVaugh 1983). 

The generic name Dasyochloa Willd. first appeared in Steudel (1840) as a 
synonym of Uralepis. Two species of Dasyochloa were listed, D. avenacea 
Willd. and D. pulchella Willd., both as "nomina nuda." Although Rydberg 
(1906) validated Dasyochloa with an English description as part of a key, it 
has been regarded as a synonym o'i Erioneuron. Recently Caro (1981) trans- 
ferred the species of the genus Erioneuron (excluding E. pilosum) to Dasyochloa, 
based on the lemma morphology. Sanchez (1983) reported that the ana- 
tomical features of Dasyochloa formed the most important difference within 

Erioneuron s.l. She recognized Dasyochloa as a monotypic North American 
genus, consisting oi D . pulchella . 

Leaf blade anatomy of North American species o(Erioneuron has not been 
fully studied. This study analyzes the leaf anatomy oi Erioneuron and com- 
pares the results with the studies from Argentina. The results of this analysis 
will add to the anatomical characteristics of the leaf epidermis not reported 
by Sanchez (1983). 

Leaf blades were selected from population samples from across the geo- 
graphic range of each species. Table 1 lists the specimens analyzed. At least 
three leaf blades were selected from each specimen so that leaf cross-sec- 
tions and adaxial and abaxial epidermes could be observed and compared. 
Leaf blade sections one cm long were taken from the specimens. Leaf blades 
collected in the field were fixed in FAA for 24 hours and then transferred to 
70% ethanol. Leaf blades from dried specimens were placed in a high mo- 
lecular weight solution of 1:3, photo-flo 200 and water until thoroughly 
imbibed, and then transferred to 70% ethanol. Leaf blades for cross-section 
analysis were embedded in paraffin using the standard techniques (Berlyn 
& Miksche 1976). Fifteen sections per plant specimen were sectioned with 
a rotary microtome at 10-15 pm thickness, stained with safranin and fast 
green, and made permanent with Permount. 

Adaxial and abaxial leaf epidermes were studied following rehydration 
in a high molecular weight solution. Leaf segments were then placed in a 
clorox solution for a few minutes to bleach the chlorophyll, placed flat on a 
microscope slide and scraped with a razor blade until the epidermis, meso- 
phyll, and vascular bundles were removed. The remaining epidermis was 
stained with Azo-Black, washed with drops of 90% ethanol, and made 
permanent using Euparal and ethanol. In some instances only small frag- 
ments of the epidermis could be obtained because of the furrowed leaf blades 
and sclerenchyma on the margins. In such cases the remaining epidermis 
was not stained to avoid the possibility of losing the tissue in the destain- 
ing process. The silica cells, silica bodies, microhairs, macrohairs, and prickle 
hair variations were recorded for each taxon. The descriptions of leaf 
transections and epidermal structures follow the terminology used by 
Metcalfe (I960) and Ellis (1976, 1979). All drawings are original and were 
made with the aid of a drawing tube. 

Examination of the leaf blade oi Erioneuron confirms that this genus has 
the Clondoid leaf anatomy as described by Brown (1958), Metcalfe (I960), 

E. nealleyi (Vasey) Tateoka 

MEXICO: Coahuila: Arceaga, Hatch et al. 5i 
Parras, Valde's-R. 1576. UNITED STATES. ' 


Dasyochloa pulchella (H.B.K.) Willd. ex Rydb. 

MEXICO. Coahuila: Sakillo, Valde's-R. 1532, Hatch et al. 5055a. San Luis Potosi: Guadalcazar, 
Valde's-R. 1642, 20 mi N of San Luis Potosi, Valde's-R. 1702. UNITED STATES. New Mexico. 
Grant Co.: Valde's-R. 1683, Otero Co.: Morden 672. Texas. Presidio Co.: Valde's-R. 1686, 1691. 

Tateoka (1961), Sanchez (1979a), and Renvoize (1983). Each bundle sheath, 
with its associated radial chlorenchyma, constitutes a discrete structural 
unit separated from similar adjacent units by large, clear bulliform cells. 
The radial chlorenchyma cells form one layer and are long, narrow, radially 
arranged, and contain few chloroplasts. This anatomical structure indicates 
that these grasses belong to the "C4 plants" or "Kranz PS" as described by 
Brown (1977). The leaf epidermis of these grasses corresponds to Renvoize's 
(1983) general description of the Eragrostideae. Complete description of 
each species is given in Valdes-Reyna (1 985). 

Erionhuron Nash 
Laminas are V-shaped, i.e. conduplicate, narrow to standard, the adaxial 
side slightly sinuous with ribs and furrows present. The abaxial side of the 
lamina is sinuous with well-marked ribs and furrows present, containing a 
prominent central vein. Sclerenchyma are associated with the vascular bun- 
dles (VB's), and a small strand sometimes forms a girder between the bundle 
and the abaxial and adaxial epidermis. Bulliform cells are inflated and fan- 
shaped. The leaf epidermis contains intercostal long cells on both epidermes 
that are moderate to strongly sinuous. Stomata occur in 1-2 rows per inter- 
costal zone with triangular subsidiary cells. Intercostal short cells occur as 
silico-suberose pairs between long cells. One papillae per cell occurs on the 
adaxial surface. Prickle hairs are distributed on intercostal and costal zones. 

Microhairs have a spherical distal cell. The macrohairs are unicellular. The 
costal zone has a row of short cells, each alternating with silica cells. Silica 
cells are mostly dumbbell- to saddle-shaped. 

Erioneuron pilosum (Buckl.) Nash 

Transverse section (Figs. 1 & 5). — Lamina is V-shaped, with a narrow to 
standard angle, 2.4 mm wide, 0.12 mm thick, 13-15 vascular bundles 
wide. The adaxial side is slightly sinuous with furrows and the abaxial side 
with slight to shallow furrows, the ribs well-marked. The prominent 
midvein has a well-developed keel, and the colorless cells, buUiform cells, 
and sclerenchyma cells are associated with the median vein. The buUiform 
cells penetrate through the abaxial epidermis. Sclerenchyma are absent 
adaxially and very abundant abaxially. One primary VB comprises the keel. 
The position of the VB is at the same level for all bundles of different 
orders. These bundles are situated in between the center and the adaxial 
surface. Three primary, (including the midvein), two secondary, and 10 
tertiary VB's are present, and round in outline. Mestome sheath cells are 
small, thick-walled, and surround the primary and secondary VB's, inter- 
rupting the tertiary VB's. Parenchyma sheath cells are large and thin-walled 
in the primary VB's and continuous or sometimes interrupted by scleren- 
chyma. The sclerenchyma that is associated with primary VB's forms a 
girder between the bundles and abaxial epidermis, and sometimes the adaxial 
epidermis. Secondary and tertiary VB's have adaxial sclerenchyma present 
as a minute strand consisting of a few subepidermal fibers. Sclerenchyma 
cell walls are very thick; with a well-developed sclerenchyma cap present 
on the margins of the leaf. This sclerenchyma cap is not in contact with the 
lateral bundle, and is wider than tertiary VB's. The chlorenchyma are radi- 
ate, one cell layer thick, in tabular arrangement, and interrupted by the 
sclerenchyma where associated with the bundle. The chlorenchyma of suc- 
cessive VB's are separated by buUiform cells and colorless cells. The color- 
less cells form a girder-like extension to the opposite epidermis and are 
associated with the buUiform cells. The colorless cells are smaller than 
buUiform cells, not inflated, and have one extension from each group. Across 
most of the blade, 3-6 buUiform cells occur in the furrows, and are exten- 
sive over midvein and in the adjacent furrow. The epidermal cell walls are 
thick and covered by a distinct cuticle that is continuous over the epider- 
mal cells. Prickle hairs are present. Macrohairs are absent. Papillae are present 
on adaxial surface and restricted to the intercostal zone. There is one pa- 
pilla per cell that is relatively broad, but not much more than half the 
width of the epidermal cells. 

Abaxial epidermis (Fig. 9b). — The intercostal long cells have margins that 
are deeply undulating, strongly corrugated, 50-170 pm long, and about 

:ions of Ertomuron and Dasyochloa. Blackenec 
E.pHosum, Valdes 1302. Fig. 2. E. avenaceum 
aides 5050. Fig. 4. D. pulchella, Valdes 1686. 

10 pm wide. Scomata are in 1-2 rows per intercostal zone with triangular 
subsidiary cells. Intercostal short cells are solitary or paired between long 
cells; when paired the cells are silico-suberose, with tall and narrow smooth 

attached to short cells, 20-25 pm long. The proximal cell is about 10 pm 
long, and the inflated and rounded distal cell is about 15 pm long. Prickle 
hairs are absent. Macrohairs are present on the intercostal zone. Costal zones 
have a row of paired short cells that are silico-suberosed and separated by 
normal costal short cells. These short cells have sinuous walls. Silica bodies, 
8_1() i^im long, are mostly dumbbell-and saddle-shaped, and relatively short. 
Adctxial epidermis (Fig. 9a).— Epidermis is similar to the abaxial surface 
except for the following: intercostal zone long cells are slightly to moder- 
ately undulating; dome-shaped, inflated papillae are present, one per cell, 
with unthickened walls. Prickle hairs are infrequent on the intercostal zone. 

Erioneuron avenaceum (H.B.K.) Tateoka 

Transverse section (Figs. 2 & 6). — Lamina is V-shaped with a narrow to 
standard angle, 2.2 mm wide, 0.l4 mm thick, 12-16 VB's wide. The adaxial 
side has ribs with slight to medium furrows, less than a quarter of the leaf 
thickness. The abaxial ribs are the same size to taller than the adaxial, 
appearing moniliform. The prominent midvein has a keel with one VB 
comprising the keel. The colorless cells, buUiform cells, and sclerenchyma 
cells are associated with the midvein. The buUiform cells in the adaxial 
epidermis are located above the median bundle. Sclerenchyma are absent 
adaxially and abundant abaxially. The position of the VB's is the same for 
all bundles of different orders. All bundles are positioned midway between 
the adaxial and abaxial surface. Five primary bundles (including the midvein) 
and 10 secondary bundles are present. Tertiary VB's were absent. Primary 

secondary bundles are circular in outline. Mestome sheaths are composed 

VB's. The parenchyma sheath, of large thin-walled cells, is continuous or 
sometimes sclerenchyma interrupts the primary VB. Sclerenchyma are as- 
sociated with the VB's on both the abaxial and adaxial surface and the 
strands consist of a few subepidermal fibers, sometimes forming a girder- 
like structure between the bundle and abaxial epidermis. The sclerenchyma 
cell walls are thick, with a sclerenchyma cap on the leaf margins. The chlo- 
renchyma are radiate, in tabular arrangement, and separated from succes- 
sive VB's by the buUiform cells and colorless cells. Colorless cells form a 
girder-like extension to the opposite epidermis, and are associated with 
buUiform cells but smaller. These cells are not inflated, and form one ex- 
tension from each group. The furrows have 2-4 buUiform cells and are 

Valdes-Reyna and Hatch, Anatomy of Eroneuron and Dasyochloa 421 

extensive over the midvein and in the adjacent furrows. The bulliform cells 
are inflated and fan-shaped. The cuticle is continuous over the epidermal 
cells. Prickle hairs are present on the adaxial surface. Macrohairs are ab- 
sent. Papillae are present on the adaxial surface but restricted to the inter- 

Abaxial and adaxial epidertms (Figs. 10a & 10b). — The anatomy of both 
surfaces is not different from that of £. pilosum epidermal surfaces; except 
that stomata, when seen, form one row per intercostal zone. 

Erioneuron grandiflorum (Vasey) Tateoka 

Significant differences were not observed between this species and E. 
avenaceum. iVletcalfe (I960) reported the leaf anatomical features of £. 
grandtflorum [syn. Tridens grandiflorus (Vasey) Woot. & Standi.} and they 
correspond to our description of £. avenaceum. 

Erioneuron nealleyi (Vasey) Tateoka 

Transverse section (Figs. 3 & 7). — This species is similar to E. avenaceum 
and E. pilosum except that the lamina is 20-25 VB's wide. The adaxial 
surface is slightly sinuous, and the abaxial surface is moderately to shal- 
lowly ribbed and furrowed. 

Abaxial epidermis (Fig. 1 lb).— Epidermis of £. nealleyt is similar to E. 
avenaceum and E. pilosu?n except that the intercostal long cell margins are 
moderately undulating. 

Adaxial epidermis (Fig. 11a). — Similar to the species mentioned, except 
that the prickle hairs are more abundant. 

Dasyochloa pulchella (H.B.K.) Willd. ex Rydb. 

Transverse section (Figs. 4 & 8). — Lamina is U-shaped, without a definite 
angle formed with the midrib, 0.6-0.9 mm wide, 0.1 mm thick, 7 VB's 
wide. The abaxial and adaxial longitudinal ribs and furrows are monili- 
form. The bulliform cells are fan-shaped, the central one relatively small, 
and not much larger than bundle sheath parenchyma cells. Sclerenchyma 
are associated with the VB's on the adaxial epidermis with a well-devel- 
oped strand that follows the shape of the adaxial rib. The well-developed 
sclerenchyma strand on the abaxial epidermis is wider than deep. The po- 
sition of the VB's is at the same level for all orders. The three primary 
bundles and four secondary bundles are situated midway between the abaxial 
and adaxial surface. These types of bundles have a round outline. The thin- 
walled cells of the mestome sheath surround the VB's. The parenchyma 
sheath, of large thin-walled cells, is continuous or interrupted by scleren- 

Figs. 9-12. Detail of adaxial and abaxial surface of leaf epidermis ofEriomuron and DasyocMoa 
Fig. 9. E.pi/osum, Valdh 1633. Fig. 10. E. avenaceum, Valdes 1623. Fig. 11. E. nealhyi 
Hatch and Valdes 5030. Fig. 12. D.pulchella, Valdes 1352, Morden 672. 

Valdes-Rf.yna and Hatch, Anatomy of Eroneuron and Dasyochloa 423 

chyma on the primary VB's. The sclerenchyma are associated with the pri- 
mary VB's forming a girder between the bundle and abaxial epidermis, and 
sometimes the adaxial epidermis. In the secondary VB's of the adaxial and 
abaxial epidermis the sclerenchyma are present as a minute strand consist- 
ing of few subepidermal fibers. The sclerenchyma cell walls are thick, and 
the lumen almost excluded. The well-developed sclerenchyma cap is present 
on the margin of the leaf, but not in contact with the lateral bundle. Chlo- 
renchyma cells are radially arranged around the VB's in one cell layer, and 
interrupted by sclerenchyma when associated with the bundle. The chlo- 
renchyma of successive VB's is separated by colorless cells and buUiform cells. 
Colorless cells form a girder-like extension to the opposite epidermis and 
are associated with and smaller than the bulliform cells. These cells are not 
inflated. Bulliform cells form a girder with associated colorless cells. The 
cuticle of epidermal cell walls is thickened and the associated cell wall 
occupies less than half of the depth of the cells. Prickle hairs present on both 
epidermes. Macrohairs are present. Papillae are absent on both epidermes. 

Abaxial epidermis (Fig. 12b). — Intercostal long cells are 60-150 jam long, 
IS about 10 |im wide, with deeply undulating margins. Stomata occur in 1 
row per intercostal zone, and are dome-shaped, with rounded subsidiary 
cells. Intercostal short cells are solitary or paired and situated between long 
cells. Papillae are absent. Bicellular microhairs are 20-24 pm long and 
occur over intercostal long cells in 1 or 2 rows. The proximal cell is about 
10 pm long, and the distal cell is about 12 pm long, and appears inflated 
and rounded. Prickle hairs, attached to short cells, are medium sized with 
the base as long as the stomata. They occur over the intercostal and costal 
zones. Macrohairs are present. Costal zones have a row of short cells with 
sinuous walls. Silica bodies are mostly dumbbell-shaped. 

Adaxial epidermis (Fig. 12a).— This epidermis is similar to abaxial sur- 
face except for the following: The intercostal zone cells are slightly undu- 
lating; prickle and macrohairs are abundant. 

The descriptions of £. pilosimi, E. avenaceum, and D. pukhella correspond 
with the anatomical features presented by Sanchez (1979a, 1983) and Caceres 

vs present. The abaxial ribs in Erioneun 
rger on the adaxial surface depicting 
)asyochloa pukhella has U-shaped lamii 
ws that form the moniliform s 
a in the leaf appears to be 
ships. The sclerenchyma; 

424 SiDA 16(3) 1995 

the VB's sometimes forms a girder between the bundle and the abaxial 
and/or adaxial epidermis, but generally the girder is represented by small 
strands. Dasyochloa pulchella differs from Erwnei/ron by having well-devel- 
oped sclerenchyma girders on the abaxial/adaxial epidermis. The scleren- 
chyma associated with the primary vascular bundle comprising the keel 
have been a key taxonomic character for the separation of £. avenaceum and 
E.pilosum (Sanchez 1979)- Erioneuron pHosum has a prominent midvein with 
sclerenchyma occupying more than half of the keel, whereas E. avenaceum 
has one third or less sclerenchyma occupying the midvein. 

The buUiform cells o^ Erioneuron species are fan-shaped as in D. pulchella, 
except the central one is smaller. Macrohairs are usually common on the 
adaxial epidermis of both genera. However, as mentioned by Sanchez (1983), 
D. pulchella macrohairs are more abundant on both epidermes. 

Leaf surfaces . — Features of the abaxial and adaxial epidermis are similar 
in most characters between the two genera. In Erioneuron, papillae were 
present only on adaxial surface, whereas in D. pulchella, they were not ob- 

Macrohairs are present in all species. However, they are more abundant 
in E. nealleyi on the adaxial surface, and in D. pulchella they occur on both 

A summary of the differences of the anatomical characters are presented 
in Table 2. The blades of £. avenaceum and E. grandiflorum had no differ- 
ences in the anatomy. Dasyochloa pulchella has characteristics that differ from 
all Erioneuron. Therefore, these data support the recognition oi Dasyochloa 
and Erioneuron as anatomically distinct genera. 


' D. Jones and J.K. Wipff (TABS) for their review and 
nanuscript. This is Technical Bulletin TA 31740, Texas 

/\nton, a.m. 1977. Notas criticas sobre gramineas de Argentina. II El g 

en Argentina. Kurtziana 10:57-67. 
Berlyn, G.P. and J. p. Miksche. 1976. Botanical microtechnique and cytoc 

Caceres, M.R. 1950. Los caracteres anatomicos tohares de Munroa mendoana y BLephartdacbne 

benthamiana. Rev. Argent. Agron. 17:233-240. 
Caro, J. a. 1981. Rehabilitacion del genero Dasyochloa (Gramineae). Dominguezia 2:1- 

CoRRELL, D.S. and M.C. Johnston. 1970. Manual of the vascular plants of Texas. Rennet, 

Texas: Texas Research Foundation. 
Ellis, R.P. 1976. A procedure for standardizing comparative leaf anatomy in the Poaceae 

I.: the leaf as viewed in transverse section. Bothalia 12:65-109. 
. 1979. A procedure for standardizing comparative leaf anatomy in the Poaceae 

II.: the epidermis as seen in surface view. Bothalia 12:641-671. 
Gould, F.W. 1968. Grass systematics. New York: McGraw-Hill. 

1975. Grasses of Texas. College Station, Texas: Texas A&M Univ. Press. 

1979. A key to the genera of Mexican grasses. Tex. Agr. Exp. Sta. MP- 

1422. College Station, TX. 
McVaugh, R. 1 983. Flora Novo-Galiciana: A descriptive account of the vascular plants of 

Western Mexico. Ann Arbor: The Univ. of Michigan Press. Vol. 14:436. 
Metcalee, C.R. I960. Anatomy of the monocotyledons, I. Gramineae. Oxford: Clarendon 

Nicora, E.g. 1973. Novedades agrostologicas patagonicas. Darwiniana 18:80-106, f 1-6. 
Parodi, L.R. 1934. Las grami'neas del genero Munroa. Rev. Mus. La Plata 34:171-193. 

Renvoize, S.A. 1983. A survey of leaf-blade anatomy in grasses IV. Eragrostideae. Kew 

1979b. Anatomia foliar de las especies y vanedades arge 
m. et Schult. y Enomuron Nash (Gramineae-Eragrosr 
22:159-175, f. 1-4. 
1981 . DesarroUo de la estructura Kranz en tallos de Gramir 

leae.LiUoa 35:37^0. 

1983. Dasyochloa Willdenow ex Rydberg (Poaceae), ge 
ca.Lilloa 36:131-138. 

■nero monotfpico de 

JDHL,E.T. 1840. Norn 
tTubingae. pp. 731, 
EOKA,T. 1961. A. bic 


Connor, S. 1994. New England Natives. (ISBN 0-674-61350-3, hbk.). 
Harvard University Press, 79 Garden Street, Cambridge, MA 02138- 
9983. $39.95. 274 pp. 

plants of New England ... by describing the'lvide'range oru.scITrmgTlfen^e'ras't! 

Chapter 3, "Trees in the Marketplace and in the Garden," considers uses and decorative 
value of trees. And chapter 4, "The New Yankee Forest," discusses ecology and more uses. 

white illustrations (mostly photographs but also reproductions of old illustrations related 

ing significance of wood in the history of the Republic" and "the use of the forest as more 
than mere scenery." It would be a fine addition to the list of "suggested readings" for any 
general botany or biology course or course on economic botany. And it is good reading for 
just about anybody interested in plants and peopie.^ohn W. Thieret. 



Department of Biology 

New Mexico State University 

Las Cruces, NM 88003-0001, U.S.A. 


Quercus basaseachicenu C. H. Muller, collected by LeSueur in 1936 and named by MuUer 

in 1938, was not recollected until 1985. Since that time this white oak (sect. Qiiercm) has 

J Desde entonces 
n la Sierra Madre 

dos Se presentan 
hojas para indicar 
aseaihuemt^ C. H. 

Harde LeSueur, from the University of Texas, was the first botanist to 
collect at what is presently the Parque Nacional "Cascada de Basaseachic" 
on the west slope of the Sierra iVladre Occidental in southwestern Chihua- 
hua, approximately 300 air km. W of Cd. Chihuahua. Among the exten- 
sive collections he made in 1936 was a peculiar white oak, which C. H. 
Muller named (Quercus hasaseachtcensis in 1938. LeSeuer's specimens were 
vegetative and Mullet's new species, therefore, necessarily lacked descrip- 
tions of flowers and fruit. Nevertheless, on leaf morphology alone Muller 
placed the new species in the subgenus Leucobalanus Engelm. (= subg. 
Quercus) and, with excellent insight, in the series Reticulatae Trelease. Muller 

SiDA 16(3): 427 -437. 1995 

4^« SiDA 16(3) 1995 

noted that a study of the fruit would probably remove the species from this 
section, to which he felt it was only distantly related. He also noted that 
"at present there exists no other series to which the species could be re- 
ferred, and Its characters are not sufficiently plam to serve as the basis of a 
new series." With no additional information, Camus (1938-39) and Martinez 
(1956) followed Mullet's classification of Q. basaseachtcensis exactly. 

Nearly 50 years passed before this oak was collected again. In 1985, 
during general floristic collecting at Basaseachic, and in subsequent years 
of botanical exploration in the northern Sierra Madre Occidental, I and 
associated collectors discovered about 20 plants referable to Q. hasaseachtcemis 
(Fig. 1 and Appendix). A few of these were in flower or in fruit. This paper, 
then, reports upon these collections, provides a description of habit of the 
plant and its flowers and fruits, and provides morphological and ecological 
evidence that Q. basaseachtcensis is a rarely formed hybrid, which at 
Basaseachic at least shows some segregration or backcrossing to the paren- 
tal types. A plant closely resembling the isotype was determined to have 
2n = 2A chromosomes (Rodriguez & Spellenberg 1992), the usual number 
in Quercus. 

The parental species are believed to be the extensively rhizomatous Q. 
depressipes Trek, a patch-forming low shrub, and the robust shrub or tree Q. 
rugosa Nee, two distantly related white oaks placed in different series 
(Trelease 1924; Camus 1938-39; Martinez 1956). This paper follows Nixon's 
(1993) infrageneric classification. He also notes that those oaks such as Q. 
depressipes, which are from Mexico and the southwestern United States and 
have connate cotyledons, form the Glaucoideae and probably are distinct at 
the subsectional level from those with free cotyledons, but a classification 
has yet to be devised. The hybrid between these very different white oaks 
that is under consideration here has not been observed to form self-per- 
petuating populations, and for this reason the hybrid name Q. xbasaseacht- 
censis C. H. Muller, /;rr; sp., is proposed (basionym: Q. basaseachicensis C. H. 
MuUer. 1938. Amer. Midi. Naturalist 19:582; Type: Mexico: Chihuahua, 
Cascada de Basaseachic, 6 Jul 1936, LeSueur 549; holotype: Mullet's pets, 
herb., transferred to BH; isotype: TEX! [photo at NMC}). 


Shrubs 1-1.8 m tall, few to many stemmed, rarely rather extensively 
rhizomatous and forming patches up to ca. 5 m across. Staminate aments 
15-42 mm long, with 4-20 flowers in the distal 80%, sparingly stellate 
tomentose; perianth sparingly to rather densely tomentose, 1.5 mm wide, 
about as long; anthers 3-7, glabrous, tan or reddish brown, 1.0-1.5 mm 
long. Pistillate flowers 1-4, in distal 1/3 of sparingly stellate pubescent to 
glabrate peduncle 15-41 mm long, usually only l(-3) maturing. Cups 

larina. Sire 4, Dgo., Mcpic 
Mcpio. Guerrero, 14.4 roai 
ifCd. Chihuahua, given fo 

10-12 mm wide, 8-10 mm deep, the mature so 
town densely appressed-pubescent bases, narrowed 
dish or tan, round or more or less acute tips that are 
rins. Acorns ovoid, 11-12 mm long, 8-10 mm wi 
t 1/2 included. Cotyledons pale pink, fused by the 
le center in the basal 2/3. 

Oaks are notorious for the frequency of hybridization within subgenera 
(see, for example, Stebbins 1950, pp. 61-66). Evidence for the hybrid na- 
ture o'i Quercus 'Xbasaseachkemis is provided by the habitat and by macro- 
and micromorphological intermediacy. The hybrid is always uncommon. 
Presently Q. xbasaseachkensis is known from five sites (Fig. 1). In Sites 1^ 
both putative parents are immediately sympatric; at Site 5 Q. y~basaseachicensis 

430 SiDA 16(3) 1995 

grew immediately beneath Q. n/f^osa, but the nearest Q. depressipes (along 
the highway) was noted at ca. 1.5 km distant. 

Habitat. — Quercus depressipes and Q. rugosa have about the same elevational 
range in Chihuahua and northern Durango, that is, ca. 1950-2600 m, the 
latter presently known to extend somewhat higher than the former. Quercus 
xbasaseachtcensis occurs between 1980 and 2225 m. Quercus rugosa com- 
monly occupies sites seeming to be more mesic, commonly in canyons, on 
north- or northeast-facing slopes, and often on deeper soils. Quercus 
depressipes commonly occurs on open sites, often on thin rocky soils. 

As is virtually common knowledge, and as reviewed by Grant (1981, p. 
199 ff.), hybrid plants are particularly common in disturbed areas. At 
Basaseachic (Site 1 in Fig. 1)12 putative hybrid plants have been located, 
all along the trail leading from the parking lot to the top of the falls. In 
this area, Q. rugosa is common, Q. depressipes is rare. At Basaseachic Q. 
depressipes is common on open rocky slopes with little tree cover, but in the 
forested area where Q. basaseachicensis plants occur Q. depressipes is not fre- 
quent. It may have been more common here in the past, and is now suc- 
cumbing to succession after fire (pines in the area are ca.l25 years old or 
younger, and bases of many large Cupressus are fire-scarred). Where it oc- 
curs in the Sierra Madre, Q. depressipes may increase very rapidly in open 
areas generated by fire (R. Corral D., pers. comm). One intermediate at 
Basaseachic, a few-stemmed shrub about 1.2 m tall near the top of the 
stairs where the trail crosses the Rfo Basaseachic, very closely resembles the 
isotype (TEX) (Fig. 2) in macromorphology and in the characteristics of 
the pubescence on the abaxial surface of the leaf (Figs. 4, 6) (in the various 
collections made during this study, this plant has been designated plant 
"#2"). No plants referable to Q. xbasaseachicensis have been found in other 
areas of the park even after 10 years of general collecting there to detail the 
flora. At Basaseachic the range of variation in hybrid plants suggests that 
some are either backcrosses to the parental types, or are later-generation 
segregates (Fig. 2); at the other sites only more or less exact intermdeiates 
(Fis?) are present. All of the plants at Basaseachic are few-stemmed shrubs; 
none are extensively rhizomatous. 

At the northernmost site (Fig. 1, Site 2) two plants referable to Q. 
x basaseachicensis were found where both Q. depressipes and Q. rugosa are com- 
mon. Intermediate plants occurred only near the edges of a dirt road. Both 
intermediate plants were few-stemmed non-rhizomatous shrubs. 

In the Sierra Catarina (Fig. 1, Site 3), six plants referable to Q. 
X basaseachicensis occur along a sharply defined contact between Q. depressipes, 
which forms an extensive and continuous patch across the open east face of 
a hillside, and Q. rugosa, which is the dominant tree (also shrub) on the 
north-facing slopes of a narrow canyon (cf. Boecklen & Spellenberg 1990 


89/9-6). Top row, I 
{mm 11 834). Midd\ 
2 {possibly from typ 

;s from Quercus depressipes, Q. rugosa, and putative hybrids, 
ection number; number following hyphen signifies individual 
s indicated in voucher specimens). See Appendix for locations, 
^es at left, Q. depressipes (3 at left from 9589, right leaf from 
oup of three leaves, Q. rugosa (2 at left from 10038, right leaf 
. xbasaseachkensisfromBasaseachicd-r, 8919-8, 8919-6, 9341- 
1 0042-4, LeSueur 549 iisotype,TEX], 10042-9, 8974-1, 8989- 

:he Sie 


, 9739-: 

xtensive description of this site). Intermediate plants do not 
t this line of contact. One of these intermediates is extensively 
, forming a thick patch several meters in diameter. Another 
four or five intermediate plants occur in a canyon ca. 1/2 km to the north- 
east where Q. rugosa and Q. depressipes are intermixed. 

Macro- and Mkromorphological Comparisons. — Hybrids in oaks are com- 
monly detected by macromorphological intermediacy, particularly by char- 
acteristics of the leaves (e.g., Bartlett 1951; Benson, Phillips & Wilder 
1967; Cottam, Tucker & Santamour 1982; Hardin 1975; Stebbins, Matzke 
& Epling 1947; Tucker 1961). In Table 1 several macro- and 
micromorphological features of Quercus depressipes, Q. rugosa, and Q. 
xbasaseachicensis are compared. Intermediacy of putative hybrids between 
Q. rugosa and Q. depressipes in leaf size and shape was demonstrated by mul- 
tivariate methods for plants in the Sierra Catarina (Site 3) (Boecklen & 
Spellenberg 1990), and is evident for two populations in Fig. 2. 

Even a casual examination of the abaxial surface of the leaf reveals con- 

spicuous differences between the species, and intermediacy in many of the 
characters by Quercus xbasaseachkensis . With a dissecting microscope Quercus 
rugosa is seen to be prominently buUate and minutely papillate (due to the 
prominently convex surfaces of the epidermal cells), Q. y^basaseachicensis 
less so, and Q. depressipes not at all buUate, and with the cell surfaces only 
obscurely convex. The bullate vs. non-bullate nature of the abaxial surface 
is evident in Figs. 3-5, as are the differences m the convexity of the cells. 

Several authors have shown that characteristics of the indumentum are 
important in distinguishing oak taxa (e.g., Hardin 1979; Manos 1993; 
Nixon & Steele 1981; Thomson & Mohlenbrock 1979) and useful in the 
detection of hybrids (Cottam, Tucker & Santamour (1982). To more criti- 
cally examine and illustrate features of the indumentum of the abaxial leaf 
surface, specimens were examined with scanning electron microscopes 
(ETEC Autoscan 500 at University of New Mexico; Philips 501B at New 
Mexico State University). Fresh leaf tissue was fixed in FAA, dehydrated, 
critical point dried, sputter-coated with gold, and photographed at 80x. 
Material from the isotype was treated in the same manner. 

Quercus rugosa has stellate hairs on the abaxial leaf surface (Fig. 3), whereas 
Q. depressipes does not (Fig. 5). The isotype of Q. xbasaseachicensis also lacks 
stellate hairs (Fig. 6), as do most of the plants believed to be hybrids at 
Basaseachic (Fig. 4). In the Sierra Catarina (Site 3) some intermediates have 
stellate hairs, others do not. Intermediate plants from sites 2, 4, and 5 all 
lack stellate hairs. 

The species and the intermediates, including the isotype, also differ in 
the size of a second type of foliar trichome, the small, gold or white vermi- 
form hairs on the abaxial surface of the leaf. Those of Quercus rugosa are 
large, easily seen, and often coalesce in age into dark droplets. The hairs are 
sufficiently small in Q. depressipes that even under a hand lens the leaf must 
be well lit for one to detect their presence. The hairs differ in length and 
diameter between Q. rugosa and Q. depressipes (Table 1), with those of Q. 
xbasaseachicensis being intermediate. 

The comparison of very different kinds of characters made in Table 1 
differs little from the character count procedure described by Wilson (1992) 
and indicates that Q. xbasaseachicensis probably is, in fact, a hybrid between 
the very different white oaks Q. depressipes and Q. rugosa. Hybrid plants are 
rare and always occur with one, and usually both, the putative parents. 
Based on leaf characteristics, including features of the indumentum, the 
plant in Figure 4 (plant #2 in the various Basaseachic collections we have 
made) may actually be the type plant from which LeSueur collected in 
1936. Admittedly, on habit and leaf characteristics it would be difficult at 

plant 2), showing only middle-size vermiform hairs. Fig. 5. (^mnus depressipes (#11839), 
showing only minute vermiform hairs. Fig, 6. Quercus xhasaseachkensis {LeSmur 549, TEX, 

SiDA 16(3) 1995 

f Qiiem/s depressipes. Q. 

liminate another common white oak, Q. anzonica Sarg., which 
Basaseachic sympatncally with Q. rugosa, as a parent instead of 
Characteristics of the pistillate inflorescence (Fig. 7) and the coty- 
•ovide insight. Qi/ercus xhasaseachicensis has rather long peduncles 
stillate inflorescence (Fig. 7), generally longer than those of Q. 
and shorter than those of Q. rugosa (Table 1). In Q. arizonka the 

Spellenberg, The hybri 



nun innui 



f^- x^^J 


I ' 

'W;- • 



M^? '^^ 


i of Q»ercus rugosa (10038) (left), Q. 
\ (8598) (right). Inflorescence from Q. 
3m Q. depressipes is from long end. Units 

Bocoyna, S edge of village of Bocoyna, 1 

peduncle rarely exceeds 1 1 mm in length. Plant #2 at Basaseachic has only 
partially connate cotyledons, as do several plants from the Sierra Catarina 
(Site 3). In Q. arizomca and Q. depressipes cotyledons are nearly or completely 
connate; in Q. rugosa they are distinct. At Site 3 Q. xbasaseachkensis occurs 
along a precise and very narrow line of contact between Q. rugosa and Q. 
depressipes, and west of Casas Grandes (Site 2) Q. xbasaseachicensis occurs 
where Q. depressipes and Q. rugosa are intermixed. In neither case is Q. 
xhasaseachicensis closely associated with Q. arizonka. At site 4, in northern 
Durango, Q. arizonka is also present and intermixed with Q. rugosa and Q. 
depressipes. At this site parentage of the intermediate is certainly open to 


Gratitude is expressed to numerous individuals. Mr. Toutcha Lebgue of 
the Universidad Autonoma de Chihuahua and Mr. Rafael Corral of the 
Escuela Superior de Agricultura "Hermanos Escobar" in Juarez both helped 
in aspects of the field work of this project and are co-collectors on many of 
the collections. Dr Billie Turner kindly gave permission to examine frag- 
ments of the isotype at the University of Texas by SEM. Esteban Muldavin 
prepared cores of several pines at Basaseachic to determine age of trees. An 
anonymous reviewer provided valuable comments. A National Science Foun- 

dation grant (DEB-92()9109) to Drs. Bill Boeckle 

with some of the costs of field work. A NMSU Ar: 

#92-027 financed SEM work done by Hank Adams and Mark Cu 

at NMSU and Angela Welford at UNM. Considerable assistanc 

vided by the NMSU Foundation "Friends of the Herbarium" fu 

Specimens seen for this paper, wirh herbaria of deposition (CHIH = Univ. Auconoma de 
Chihuahua; ESAHE = Coiegio de Graduados, Escuela Superior Agricultura "Hermanos 
Escobar" in Juarez [now closed; possibly to be transferred to Univ. Autonoma de Cd. Juarez}). 
Collection numbers are those of Spellenberg and various associates. Sites of collections are 

SITE 1— Chihuihua Mcpio Ocampo Pirque Nicionil Cascada de Basaseichic 
108 12S0"W 2S lO'N eleN I 9S() m in issoci inon \Mth various oaks and pines 

Q basaseathni>,M^ I tS/a/n ^^9 ^ 6 i6(Tfc\' isotvpe) S-^6" 26 Apr 19S5 (BH DAV 
CHIH SAlll Ni\I( ) S"^9^ lOLtl9S6(NMC) sg/9 16 Oct 1986 (NMC) 9:>4l 
BH IBUC NMC ir\) 9-)SS 1 Au^ i9SS(( AS C IIDIR lEB NMC) 96^"^ 1 
Aug 19SS(NMC) 9660 2 Au^ 19SS (ASU ME\U NMC US) 10042 28 Oct 
1989 (ME\U UC) 1092^ 26 Sep 1991 (MEXU MO NMC NY) 11S40 17Jun 
199^ (BRIT DAV RSA) 

Q ckpresupes S469 27 Api 19S7 (MEXU) S9/9 16 Oct 1986 (NMC) 95S9 1 Aug 
1988(CAS CIIDIR lEB NMC) 964^ A 1 Aug 1988 (NMC) 70799 25Jun 1991 
(F) lis, 9 1 Jun 199^ (NMC) 

Q )iigosa S9/7 16 0ct 19S6(NMC) 9:>4l 12Sepl9S7(BH BRIT CAS lEB NMC) 
9590 1 Au^ 19SS(NMC UC US) lOOiH 28 Oct 1989 (CIIDIR MEXU NMC) 

Q basasinhttinn 9/9^ 10 Jun I9S-((11D1R DAV MEXU NMC) 

Q deple^upi^ 9/9/ 10 jun i9S^ (CIIDIR MEXU NMC) 

Q tiii^ina 9190 lOJun 19S7 (CIIDIR MEXU NMC) 
SITE :, — Chihuahua Mcpio I Zaragoza 25 km SW of Buenaventuri 1 1 km E ( 
over Sierra Catarina 107 o8'W 29 46'N elev 2290 m m association with viriou 

Q basaseaihuemis 79^6 S Feb 19S5 (DAV MEXU NMC) S9-/0 11 Nov 19S6(r 
MO) b935 l6No% 1986 (BRIT COLO DAV I NMC RM SRSC) 92/. 1 

RSA TEX) /09)6 I 2 Oct 1991 (CI I) 
Q depnssipei 795) S Feb 1 9S5 (MEXU NMC) /0060 lO Oct 19S9 ARIZ 

CAS CIIDIR MEXU NMC) S9^7 14 Nov 19S6(BH F lEB) 92/2 ISAu^ 

(MEXU NMC N^) 9"^0 2i Sep 19SS (IBUC NMC) 
Q }Ui>osa S9^9 1 1 Nov 19S6(ASU IBUG) Sy5^ i 6 Nov 19S6(COLO DAV I 

INIF NMC NY SRSC) S956 l6Nov 19S6(MEXU NMC) /00^9 ^OOct 


3543, 12 Jul 1986 (CAS, CIIDIR, MEXU, NMC) 
, 12 Jul 1986 (MEXU, NMC) 

Mcpio. Guerrero, 14.4 road km W of Tomochic 
^660, 2 Aug 1988 (ASU, MEXU, NMC, US) 

Bartlett, H. H. 195 1 . Regression of xQuercus deamit toward Quercus macrocarpa and Quenus 

muhlenbergit. Rhodora 53:249-264. 
Benson, L., E. A. Phillips, and P. A. Wilder. 1967. Evolutionary sorting of characters in a 

hybrid swarm. I. Direction of slope. Amer. J. Bot. 54:1017-1026. 
BoECKLEN, W. J. and R. Spellenberg. 1990. Structure of herbivore communities m two 

oak {Quercus spp.) hybrid zones. Oecologia 85:92-100. 
Camus, A. 1938-39- Les chenes. Monographie du genre Quercus, tome II, genre Quercus, 

sous-genre EuQuercus. Paul Lechevalier, Paris. 
CoTTAM, W. R, J. M. Tucker and R S. Santamour, Jr. 1982. Oak hybridization at the 

University of Utah State Arboretum of Utah, Pub. #1. Salt Lake City 
Grant, V. 1981. Plant speciation (2nd ed.). Columbia Univ. Press, NY. 
Hardin, J. W. 1975. Hybridization and introgression in Quercus alba. J. Arnold Arbor. 

P S. 1993. Foliar tnchome variation in Quercus section Protobalanus (Fagaceae). 

;/., M. 1956. Los encinos de Mexico, VII. Ann. Inst. Biol. Mexico 27:373-395. 
C. H. 1938. Further studies in southwestern oaks. Amer. Midi. Naturalist 19:582- 

K. C. 1993. Infrageneric classification oi Quercus (Fagaceae) and typification of 
nal names. Pp. 25-34 In: A. Kremer, P S. Savill and K. C. Sterner (eds.). Genetics 
ks, Ann. Sci. Forest. 50, Suppl. 1, 290 pp. 

and K. P Steele. 1981. A new species oi Quercus (Fagaceae) from southern 

.rnia.Madroi^o 28:210-219. 

JEZ, T, S. and R. Spellenberg. 1992. Chromosome numbers for five Chihuahuan 
;s oi Quercus (Fagaceae). Phytologia 72:40-41. 

., G. L., Jr. 1950. Variation and evolution in plants. Columbia Univ. Press, NY. 
I E. C. Epling. 1947. Hybridization in a population of 

Querelas marilandica and Quercus ilicifolia. Evolu 
Thomson, P M. and R. H. Mohlenbrock. 1979- Foliar trichomas oi Quercus subgt 

Quercus in the eastern United States. J. Arnold Arbor. 60:350-366. 
Trelease, W. 1924. The American oaks. Mem. Natl. Acad. Sci. 20:1-255 + 420 plat 
Tucker, J, M. 1961. Studies in the Quercus undulata complex. I. A preliminary statem 

Amer. J. Bot 48:202-208. 



Li, Shi you and Dent T. Adair. 1994. XI Shu. A Promising Anti-tumor 
and Anti-viral Tree for the 21st Century. (ISBN 0-938361-11-2, 
hbk.). The Tucker Center, College of Forestry, P.O. Box 6109, SFA 
Station, Nacogdoches, TX 75962, (409) 468-4600 voice; 409-468- 
1 195 fax. $45.00. 268 pp., 18 color & 6 b&w photos, 6x9. 

This book is a monograph about Xi Shu and camptochecins. It includes two parts. Part 
I includes two sections and 13 chapters of text: 1) Camptothecins: drug discovery history, 

A foreword is provided by Dr. Monroe E. Wall (Chief Scientists of the Research Triangle 
Institute, North Carolina), the discoverer of drugs camptothecin and taxol. And a preface 
is provided by Dr. Beppino C. Giovanella of Stehlin Foundation for Cancer Research. 

Hopkins, William G. 1995. Introduction to Plant Physiology. (ISBN 0- 
471-54547-3, hbk.). John Wiley & Sons, Inc., 605 Third Avenue, 
New York, NY 10158, (212) 850-6336. $75.95. 464 pp. Illustrated. 

e; 19) Measuring Time: Photoperiodism and Rhythmic Phenomena; 20) 
1 Plant Development; 21) The Physiology of Plants Under Stress; and 22) 



United States Department of Agriculture 

Systematic Botany & Mycology Laboratory 

Bldg. 003, B ARC -West 

Beltsville, MD 20705, U.S.A. 

A historical perspective and discussion of the generic relationships is presented fc 

staria. A list of accepted taxa previously placed in Paspalidium is given. Nineteen ne- 

. gausa, S. globoidea, S. spartela, S. grandtspkulata, S. maequalis, S. jubiflora, S. paludivaga, . 
tra, S. reflexa, S. rettglumts, S. scabnfolia, S. tabulata, S. uda) and one new name {Setan 

Paspaltdium y Setaria. Se concluye que las especies de Paspalidium 

clementti, S. constrkta, C. mniformts, S. gausa, S. globoidea, S. sparti 
maequalis, S. jubiflora, S. paludivaga, S. rara, S. reflexa, S. retiglumis, , 

The generic distinction between Paspalidium Stapf and Setana P. Beauv. 
has been a topic of academic interest among agrostoiogists knowledgeable 
in the phenetic relationship between these genera. Setaria includes about 
114 species (Webster 1993) concentrated in the tropics and sub-tropics of 
the world. Presence of bristles, disarticulation at the base of the spikelets, 
indurate upper floret, and a muticous upper floret are characters that sepa- 
rate Setaria from other genera of the Paniceae R. Br. (Webster 1992). 
Paspalidium includes about 28 species. It is concentrated in Australia where 
Webster (1987) recognized 23 species. The remaining five species are na- 
tive to Asia and Africa. 

Stapf (1920) separated Paspalidium from Setaria on inflorescence form 
and structure. That is, Paspalidium was characterized by racemose primary 
branches and Setana by a contracted spike-like panicle. Subsequent au- 
thors used similar characters and the presence or absence of bristles to sepa- 

440 SiDA 16(3) 1995 

Hitchcock and Chase (1910) separated Panicum subgenus Paurochaetum 
from other species olPamcum on the presence of a point or bristle extend- 
ing beyond the uppermost spikelet in subg. Paurochaetum. Pilger (1940) 
placed subgenus Paurochaetum as a section oiSetarta. Hitchcock (195 1) sepa- 
rated Setaria from Panicum on the presence of bristles forming an involucre 
in Setaria, but retained Paurochaetum in Panicum. Rominger (1962) did not 
discuss Paspalidium but placed Paurochaetum as a subgenus oi Setaria. 

Veldkamp (1980) noted that Paspalidium was probably not distinct from 
Setaria. Clayton and Renvoize (1986) did not formally place Paurochaetum. 
They recognized both Setaria and Paspalidium but indicate that intermedi- 
ate species make the distinction arbitrary. 

A complete comparative treatment of the 23 Australian Paspalidium was 
presented by Webster (1987). I stated the following: "Contrary to what the 
various flora treatments in North America, Africa, and Australia lead one 
to believe, Paspalidium cannot be distinguished from Setaria on the pres- 
ence or absence of bristles and the spikelet characteristics are essentially 
identical. The bristle to spikelet relationships in Setaria are so varied that it 
cannot be used to separate the genera." I tentatively distinguished these 
genera on the arrangement of the primary inflorescence branches. Paspalidium 
was characterized by secund or distichous primary branches, whereas in 
Setaria the primary branches could originate from any point on the main 
axis (i.e. quaquaversal). This character was applied in subsequent publica- 
tions distinguishing among the Paniceae genera (Webster 1988; Webster 
& Valdes-Reyna 1988; Webster et al. 1989; Webster 1992). 

Davidse and Pohl (1992) stated that Paspalidium is characterized by 
abaxial spikelets born in unilateral spikes. They made new combinations 
in Paspalidium for Setaria chapmanii and six West Indian species with an 
inflorescence form similar to the Australian Paspalidium. The New World 
mainland taxa with similar inflorescence characteristics to those of the 
Australian and West Indian "Paspalidiums" groups were retained in Setaria. 

Webster (1993) stated that "when the full range of variation in Setaria is 
considered, it is morphologically identical to Paspalidium." Finally, 
Veldkamp (1994) concluded that Paspalidium cannot be delimited from 
Setaria. He reduced it to Setaria, and transferred the Southeast Asian spe- 
cies to Setaria, including the type species oi Paspalidium. 

In review, all species o'i Paspalidium possess a bristle terminating a branch. 
The spikelets are arranged in a tight or loose arrangement on the branches 
depending on the number of spikelets and the length of the pedicels. In 
some species, bristles subtend all spikelets whereas in others only the ter- 
minal spikelet of a branch is subtended by a bristle. There is continuous 
variation between these forms. A wide range in variation for these charac- 
ters is also found in Setaria. As in Paspalidium, the bristle to spikelet tela- 

Webster, Nomenclatural changes in Setaria and Paspalidium 44 1 

tionship is correlated to the development of the primary branches. Species 
with elongate or pronounced primary branches frequently have some spike- 
lets that lack subtending bristles. However, a bristle will terminate the 
primary and secondary branches. Arrangement of the primary branches 
(i.e. secund, distichous, and quaquaversal) is difficult to apply and the ge- 
neric significance is open to question. Since the genera cannot be satisfacto- 
rily separated, Paspalidium is best treated as a synonym o{ Setaria. A com- 
prehensive phylogenetic study of Setana is required before a decision is 
made concerning the recognition of subgenera and sections within Setaria. 
The foUowmg is an alphabetical list of accepted species with new com- 
binations where needed. 

^ta^ia albovillosa (S.T. Blake) R.D. Webster, comb. nov. Paspalidium 
albovillosum S.T. Blake, Proc. Roy. Soc. Queensland 62:96. 1952. Type: Blake 10947 
(holotype: BRI!; isotype: CANB!). 

Paspalidium radiatum var. hirsutum Vickery, Concr. New South Wales Natl. Herb, 1 :334. 
1951. Type: CO. Cross NSW no. 8944 (holotype: NSW!). 

Distribution. — Australia: woodlands of Queensland and New South Wales. 

etaria aversa (Vickery) R.D. Webster, comb. nov. Paspalidium aversum Vickery, 
Contr. New South Wales Natl. Herb. 1:3.31. 1951. Type: T.F. Mau s.n. (holotype: 
NSW no. 8992!). 
Distribution. — Australia: woodlands of Queensland and New South Wales. 

etaria basiclada (Hughes) R.D. Webster, comb. nov. Paspalidium basidadum 
Hughes, Bull. Misc. Inform. 318. 1923. Type: S toward 200 (holotype: K!). 
Distribution. — Australia: shrublands, grassland, and arid regions of West- 
-n Australia, Northern Territory, South Australia, and Queensland. 

. Paspalidium caespitosum C. E. Hubb., 

Distribution. — Australia: woodlands to and shrublands of Queensland 
nd New South Wales. 

Type: Chapman s.n. (lectotype, selected by Hitchcock & Chase 1910, US!). 
Dtstnhutwn.^'UnMed States: Florida. Mexico: Yucatan. West Indies: Cuba 
nd Bahamas. 

Setana clementii (Domm) R.D. Webster, comb. nov. Paspalidinm demmfn 
(Domin) C. E. Hubb., Bull. Misc. Inform. 447. 19.M. Pamcum dementu Domin, J. 
Linn. Soc, Bot. 41:272. 1912. Type: E. Clement s.n. (holotype: either BM or PR). 
Distribution. — Australia: arid grasslands and shrublands of Western Aus- 
tralia, Northern Territory, South Australia, Queensland, and New South Wales. 

Setaria constricta (Domin) R.D. Webster, comb. nov. Paipalidiumcomtnctum 
(Domin) C. E. Hubb., Bull. Misc. Inform. 447. 19.M. Panicum comtnctim Domin, 
Biblioth. Bot. 20(85):3()2. 1915. Pamcum flavidum van tenuis Benth., Fl. Austral. 

Distribution. — Australia: arid grasslands and shrublands of Western Aus- 
tralia, Northern Territory, South Australia, Queensland, and New South Wales. 

Paspalidium gradle (R. Br.) Hughes var. debile Vickery, Contr. New South Wales Natl. 

Herb. 1:331. 1951. Type.J.H. Camfield 5, 1901 (NSW no. 9168!). 
Distribution. — Australia: woodlands and shrublands of Queensland and 
New South Wales. 

Setaria distans (Trin.) Veldkamp, Blumea 39:376. 1994. Paspalidium distans 
(Trin.) Hughes, Bull. Misc. Inform. 317. 1923. Panicum distans Trin., Spec. Gram. 
2:t. #172. 1829, non Saltz. ex Steud. 1853. Type: R. Brown 6098 (holotype: K; 

Setaria distantiflora (A. Rich.) Pilg. in Engl. & P 
(ed. 2) I4e:72. 1940. Paspalidnm distanttflorum 
Pohl, Novon 2:106. 1992. Pamcum distantiflorum A. Ri 

Distrtbutton.—'^est Indies: Cuba and ] 

Setaria flavida (Retz.) Veldkamp, 
(Retz.) A. Camus in Lecomte, Fl. Gei 
Retz., Obs. Bot. 4:15. 1786. Type: 

Webster, Norn 

Distribution. — Australia: woodlands of Queensland and New South Wales. 

Setaria geminata (Forssk.) Veldkamp, Blumea 39:377. 1994. Paspaltdtum 
geminatum (Forssk.) Stapf, Fl. Trop. Afr. 9:585. 1920. Pamcum geminatum Forssk., Fl. 
Aegypt.-Arab. 18. 1775. Type: Forsskal s.n. (holotype: C). 
Distribution. — Native to Africa and Asia, now widely introduced in the 
tropics and subtropics of the world. 

Setaria geminata (Forssk.) Veldkamp var. paludivaga (Hitchc. & Chase) R.D. 

Webster, comb. nov. Pasl,alidtumgeminatmn{Vo^ssk.)Sz^piv2ir.paludivagum{U:vtchc. 

Ik Chase) Gould, Southw. Naturalist 15:391 • 1971. Paspalidium paludivagum (Fiitchc. 

& Chase) Parodi, Gram. Bonaer. ed. 3. 85, 89. 1939- Paniam paludivagum Hitchc. & 

Chase, Contr. U.S. Natl. Herb. 15:132. 1910. Type: Nash 746 (holotype: US!). 
Distribution.— Wet areas of United States (Florida), Mexico, Guatemala, 
and West Indies. 

Setaria globoidea (Domin) R.D. Webster, comb. nov. Paspalidmm gbboideum 
(Domin) Hughes, Bull. Misc. Inform. 317. \92i.Pankum globoideum'Domin,'R.t^en. 
Spec. Nov. Regni Veg. 10:119. 1911. Type: Wuth s.n. (lectotype: K). 
Distribution. — Australia: woodlands of Queensland and New South Wales. 

Setaria grandispiculata(B. K. Simon) R.D. Webster, comb. nov. Paspaltdtum 

grandispiculatum B. K. Simon, Austrobaileya 1:465. 1982. Type: Peart 1990 (holo- 
type: BRI!; isotypes: CANB!, K, L, MO, NSW!). 
Distribution. — Australia: woodlands of Queensland. 

Setaria inaequalis (R Muell.) R.D. Webster, comb. nov. Paspaltdtum tnaequak 
(F Muell.) Hughes, Bull. Misc. Inform. 317. 1923. Panictm tnaequak F. Muell., 
Frag. 8:189. 1874. Type: Mueller s.n. (holotype: MEL; isotype: K). 
Distribution. — Australia: tropical woodlands of Queensland. 

Setaria jubiflora (Trin.) R.D. Webster, comb. nov. Paspalidium jubiflorum 

(Trin.) Hughes, Bull. Misc. Inform. 3l7. 1923. Pamcum ptblflorum Tun., 

Gram. Pan. Diss. 2:130. 1826. Pamcum flavidtim var. jubiflorum (Trin.) Domin, 

Biblioth. Bot. 20(85):300. 1915. Type: Lindley s.n. (holotype: CGE?). 

Distribution.— Axxstt^Yi^: arid grassland to sub-humid woodlands of 

Western Australia, Northern Territory, South Australia, Queensland, and 

New South Wales. 

Setaria leonis (Ekman ex Hitchc.) Leon, Contr. Ocas. Mus, Hist. Nat. 

'. Ind. 295. 1936. Type: E/fe, 
!.— West Indies: Cuba. 

Setaria ophiticola (Hicchc. & Ekman) Leon, Contr. Ocas. Mu 
Colegio "De La Salle" 8: 1 63 . 1 946. Paspalidmm ophiticola (Hit 
Davidse & R. W. Pohl, Novon 2:106. 1992. Pankum ophiticola Hitcl" 
Hitchc, Man. Grasses W. Ind. 293. 1936. Type: Ekman 12712 (hol 

Distribution.— West \Mx 

es: Cuba. 

staria pradana (Leon ex 
Colegio "De La Salle" 
Davidse & R. W. Pohl, N 
Man. Grasses W. Ind. 294. 

Hkchc.) Leor 
8:164. 1946.; 

1, Conrr. 

Distribution. — Wes 1 1 nd 1 1 

es: Cuba. 

Setaria punctata (Burm. f.) Veldkamp, Blumea 39:381. 1994. Paspalum 
punctatum (Burm. f.) Srapf ex Ridl., Fl. Malay Penins. 5:218. 1925. Paspalidium 
punctatum (Burm. f.) A. Camus in Lecomte, Fl. Gen. Indo-Chine 7:419. 1922. Pant- 

Clayton & Renvoize 1982, BM). 

' (LECTOTYPE, Selected by 

Paspaltduim mucronatum (Roem. & Schult.) Ohwi, Acta Phytota 
Pankum mucronatum Roem. & Schult., Syst. Veg. 2:425. 181 

LX.Geobot. 11:33. 1942. 

Distribution. — Africa, Asia, and Pacific Islands. 

Setaria rara (R. Br.) R.D. Webster, comb. nov. Paspalidm 
Bull. Misc. Inform. 318. 1923. Pamcun rarum R. Br., Pro 

v..™ (R.Br.) Hughes, 

Distribution. — Australia: arid grasslands and sub-humind woodlands of 
Western Australia, Northern Territory, Queensland, and New South Wales. 

Setaria reflexa (R.D. Webster) R.D. Webster, comb. nov. Paspalidium reflexum 
R.D. Webster, Australian Paniceae 166. 1987. Type: Latz 4847 (holotype: NT). 
Distribution.— A\xstz^\i2i: arid grasslands of Western Australia, Northern 
Territory, and South Australia. 

Setaria retiglumis (Domin) R.D. Webster, comb. nov. Paspatidum retiglmm 

(Domin) Hughes, Bull. Misc. Inform. 317. 1923. Pankum nUglume Yiomin, Repert. 

Spec. Nov. Regni Veg. 10:1 19. 191 1. Type: Mueller s.n. (holotype: MEL; isotype: K). 

Distribution. — Australia: woodlands of Western Australia, Northern Ter- 

Paspalidium gracile (R. Br.) Hughes, Bull. Misc. Inform. 318. 1923 Pantcum gractle R. 
Br., Prodr. 190. 1810, non Setana gracilis Kunch (1824), nee Spreng. ex Trin. (1835). 
Type: R. Brown 6096 (holotype: K). 
Distribution. — Australia: woodlands, shrublands, and grasslands of West- 
rn Australia, Northern Territory, Queensland, and New South Wales. 

.etaria subtransiens Hitchc. & Ekman in Hitchc, Man. Grasses W. Ind. 
351. 1936. Paspalidium subtransiens (Hitchc. & Ekman) Davidse & R. W. Pohl, 
Novon 2:106. 1992. Type: Ekman 16828 (holotype: US). 
Distribution.— West Indies: Cuba. 

Distribution. — Australia: shrublands and grasslands of Western Australi; 

Setaria uda (S.T. Blake) R.D. Webster, comb. nov. Paspalidium udum S.T Blak 
Proc. Roy. Soc. Queensland 62:98. 1952, Type: 5,7: Blake 16639 (holotype: BR 
Distribution. — Australia: tropical woodlands of Northern Australia an 


Setaria utowanaea (Scribn.) Pilg. in Engl. & Prantl, Nat. Pflanzenfan 
(ed. 2) I4e:72. 1940. Paspalidium utowanaeum (Scnbn.) Davidse & R. W. Poh 
Novon 2:106. 1992, Panicum utowanaeum ^cnhrx. in Millsp., Publ. Field Columbia 
Mus., Bot. Sen 2:25. 1900. Type: Millspaugh 702 (holotype: F). 
Distribution. — West Indies: Greater and Lesser Antilles. 

xtended to Dr. John Wiersen 

)AViDSE, G. and R.W. Pohl. 1992. New taxa and nomenclatural combinations o 

Mesoamerican grasses (Poaceae). Novon 2:81-110. 
Iitchcock, A.S. and A. Chase. 1910. The North American species oi Panicum. Conn 

U.S. Natl. Herb. 15:1-332. 
iiTCHCOCK, A.S. 1951. Manual of the grasses of the United States, 2nd ed. Revised by A 

Chase. Washington, D. C: U.S. Dept. Agric. Misc. Publ. 200. 

ROMINGER, J.M. 1962. Taxonomy oi Setaria (Gramineae) in N 

Mon. 29:1-132. 
Staff, O. 1920. Gramineae. In: D. Prain (ed.), Flora of Tropic 

Reeve & Co. 
Veldkamp, J.R 1980. Setana divalis (Ridl.) Veldk. comt 

gratulatorius m honorem H.C.D. de Wit. Misc. Pap. Landbc 

19:315-320. Miscellaneous Paper 19. 

Bot. 13:576-609. 
Webster. R.D. and J. Valdes-Reyna. 1988. Genera of Mesoamerican Paniceae (Poace; 

Panicoideae).Sida 13:187-221. 
Webster, R.D., J. Kirkbride, and J. Valdes-Reyna. 1989. New World genera of t 

Paniceae (Panicoideae; Poaceae). Sida 13:393^17. 
Webster, R.D, 1992. Old World genera of the Paniceae (Poaceae: Panicoideae). Sida 1 






Herbarium, Department of Botany 
Oklahoma State University 
Stillwater, OK 74078-0289, U.S.A. 
Department of Statistics 
Oklahoma State University 
Stillwater, OK 74078, U.S.A. 

Yunck. rather than Lepia 


Engelm. as 

, previously place 

;d. Analyses of 


variation, via univariate ; 


;is, UPGM/ 

cipal component analysis, and 

discriminant analysis, in 

t that It is c 

listinct from C. t 


nd C. cusptdata 

Engelm. but morphologic 

rally s 



specially var./.. 


1 of intersp 

ecific hybridizati. 

ons reveal that 

reproductively isolated from t 

he other ta) 

<a. These data suggest that C. c 

menuata merits 

continued recogmtion as 


tinct specie. 

El examen de las relac 



rdTrara paras 

ita Cuscuta am 


especies del genero revela 



>cada en la subseci 

:. Indecorae en vi 

tz de la subsect. 

Leprdanche como se habf 

a hec 

ho previam, 

ente. Los analisis 

; de la variacic 

in morfologica, 



ntes principles 

y analisis discriminante, ii 


m&A&C. compacta 




eiante a C. / 

ndecora Choisy, esj 


iv?ir:. longisepala 

)tenidos en un pr 

laciones interespecificas revela 

que C. attenuata esta aisla 

Ida re 

productivamente de los otros 

taxa. Estos dat 

OS sugieren que 

Because of its rarity and uncertain taxonomic status, Cuscuta attenu, 
Waterf. has merited much attention (Tyrl et ah 1978; Taylor & Taylor 191 
Prather 1990; Prather & Tyrl 1993). The species was described from pla 

448 SiDA 16(3) 1995 

collected from two neighboring populations in the Red River Hoodplain of 
extreme southeastern Oklahoma (Waterfall 1971). Prior to 1989 the spe- 
cies was known from only four populations within a few km of one another 
(Tyrl et al. 1978; Taylor & Taylor 1980). Because of its limited geographi- 
cal distribution, it was considered to warrant possible designation as an 
endangered species and was declared a Category 1 species by the U. S. Fish 
& Wildhfe Service's Office of Endangered Species (1980 FR 45:82500; 1985 
FR 50:39526). Prather (1990) and Prather and Tyrl (1993) reported four 
extant and six historical populations in Kansas, Oklahoma, and Texas. As a 
result, the status of the species was modified to Category 2 (1993 FR 

Waterfall (1971) tentatively placed C. attenuata in subsect. Lepidanche 
Engelm. on the basis of his interpretation that the calyx was polysepalous. 
He suggested, however, that formation of a new subsection to accommo- 
date the species might be appropriate because its capsule shape and the 
distribution of its floral bracts were not consistent with Yuncker's circum- 
scription of subsect. Lepidanche (Yuncker 1965). Tyrl et al. (1978) echoed 
Waterfall's uncertainty about the species' relationships and called for taxo- 
nomic investigations of the species and its putative relatives. 

Among the species of subsect. Lepidanche, Waterfall stated that C. attenuata 
most closely resembled C. compacta Juss. but distinguished them on the 
basis that: (1) C. attenuata has a pedicel while C. compacta does not, (2) C. 
attenuata has only one floral bract which is at the base of the pedicel while 
C. compacta has 1-10 bracts which are situated along the length of the 
pedicel, and (3) C. attenuata has lanceolate, attenuate sepals while C. compacta 
has ovate, obtuse sepals (Table 1). Waterfall also stated that C. attenuata 
resembled C. cuspidata Engelm. in the presence of pedicels, which all other 
species in the subsect. Lepidanche lack. They are different in that C. cmpidata 
has a much more open inflorescence; ovate, cuspidate sepals; and usually 
one or two bracts along the pedicel (Table 1). 

During a preliminary examination of herbarium specimens, including 
the holotype, it was discovered that the calyx of C. attenuata is gamosepalous 
and not polysepalous as Waterfall stated in his diagnosis. Because of this 
fusion, C. attenuata clearly seems better placed within subsect. Indecorae 
Yunck. rather than in subsect. Lepidanche. The two subsections are not 
thought to be closely related though both are positioned within sect. 
Cleistogrammica Engelm. (Yuncker 1932, 1965). With the exception of its 
somewhat dense inflorescence, all of the characters of C. attenuata are better 
accomodated in subsect. Indecorae (C^dkAt 1). 

Within the subsection, C. attenuata is morphologically similar to C. 
indecora var. longisepala Yunck., and specimens of C. attenuata key to var. 

Prather et al., Taxonomic investigations of Cuscuta attenu: 
Table 1. Comparison of morphological characters o{ Cuscuta species 

longisepala in Yuncker's 1965 key. Waterfall's description of C. attenuata 
(Waterfall 1971) and Yuncker's description of C. indecora var. longisepala 
(Yuncker 1965) are similar for every character state. In 1965, Yuncker ex- 
amined one of the specimens identified in this study as C. attenuata {C.J. 
Eskew 1393, OKL). Recognizing the distinctiveness of the specimen, he 
made the following annotation: "C. indecora Choisy. The long narrow calyx 
lobes would make it var. longisepala Yunck. However, the specimen looks 
teratological and may not be the variety but only an abnormal form." 

Cuscuta indecora is widespread in North and South America and is highly 
variable throughout Its range (Hunziker 1950; Yuncker 1965;Beliz 1986). 
The infraspecific classification is somewhat controversial and Yuncker (1920, 
1932, 1965) accepted different varieties in each of his treatments. Cuscuta 
indecora var. longisepala occurs throughout much of the range of C. indecora, 
including South America where the variation is similar to that found in 
North America (Hunziker 1950). Beliz (1 986) does not mention C. indecora 
var. longisepala or list it in synonymy. Variability over the range of C. indecora 
is not discussed nor were representative specimens west of Arizona cited. 

The few chromosome numbers known for taxa of the two subsections are 
invariant. Cuscuta indecora and C. glomerata Choisy (subsect. Lepidanche) are 
both reported as n= 15 (Freeman & Brooks 1988; Pinkavaet al. 1974) while 
C. attenuata is reported as 2n = 30 (Prather & Tyrl 1993). 

An investigation of the relationship of C attenuata to other taxa oi Cuscuta 
by means of analyses of morphological variation and a program of interspe- 
cific hybridization was undertaken in an attempt to clarify the taxonomic 
position and rank of the species. 

Analym of morphological variation . — To examine morphological variation 
within and between taxa, 186 herbarium specimens oiCuscuta from DUR, 
TEX, TULS, UARK, and the herbarium at Cameron University in Lawton, 
Oklahoma were examined — 10 of C. attenuata\ 50 each of C. compacta, C. 
cuspidata, and C. indecora var. indecora; and 26 of C indecora var. longisepala 
(Prather 1990). 

In a preliminary analysis, five flowers each were examined on several 
specimens of each taxon. It was determined that variation among flowers of 
the same plant was negligible, therefore one flower was used from each 
specimen, which was treated as an OTU. One individual per population 
was examined. The data recorded for each specimen are presented in Prather 
(1990). Because Cuscuta lacks roots and well-developed leaves, and because 
stem features could not be accurately scored from herbarium specimens, 
only floral characters were used. 

One flower, with its pedicel and bract(s), was removed from each speci- 
men. To minimize variation resulting from flower age, all flowers selected 
had dehiscing anthers, a stage which lasts only a short time (Prather & Tyrl 
1993). Observations were made after softening each flower by boiling in 
water to facilitate examination of the inner floral parts (Yuncker 1920). 
Samples were examined with a dissecting microscope at a magnification of 
30x to score characters. Measurements were recorded to the nearest 0.1 
mm using an ocular micrometer. Many of the measurements were incorpo- 
rated in the analyses as ratios to minimize the effect of size which may be 
influenced by environmental factors. Forty-four characters were scored for 
each sample. The first 27 characters (Table 2) were qualitative and there- 
fore suitable for use in all analyses. Characters 28-44 (Table 3) were quali- 
tative characters and thus could only be used in the multivariate analyses. 

For characters 1 and 5-27 an unprotected LSD test was performed. Char- 
acters 2-4 were excluded because the values were invariant among indi- 
viduals of at least one taxon. The multivariate analyses comprised UPGMA 
clustering, a principal component analysis, and a discriminant analysis. 
Statistical Analysis System (SAS Institute, Inc. 1985) was employed to 
perform these analyses except the unprotected LSD tests which were per- 
formed using Statview (Abacus Concepts, Inc. 1992). The UNIVARIATE 
procedure of SAS was used to confirm that the assumption normality of the 
LSD test was not violated. The UPGMA clustering analysis was performed 
using the AVERAGE option of the CLUSTER procedure on data that had 
been standardized by the STD option which changes the mean to zero and 
the standard deviation to one. The varimax rotation method was used in 
the principal component analysis. Because the other statistical methods 

Prather et al., Taxonomic investigations of Cuscuta a 

i.3) 24-3= 


Si DA 16(3) 1995 

had established that C. wmpaUa and C tuspidata were easily distinguished 
from C. attenuata, only C attenuata and the two varieties of C indecora were 
examined in the discriminant analysis. Prior probability of the discrimi- 
nant analysis was set proportional to the number of specimens of each taxon 
used in the analysis. 

Interspecific hybridizations. — Parasitized host plants of all taxa were trans- 
ported to the plant growth facility at OSU and maintained as described 
previously (Prather & Tyrl 1993). Attempts to maintain C. compacta in the 
laboratory were unsuccessful because transplanting its woody hosts was 
not possible and cuttings did not survive under laboratory conditions. Pol- 
len of C. compacta, therefore, was collected from five individuals in the field 
and used immediately in crosses in the laboratory. Vouchers of all popula- 
tions used were deposited in OKLA. 

Individual flowers of C. attenuata were emasculated before anther dehis- 
cence and mature pollen from individuals of another taxon was manually 
transferred to the stigmas (Radford et al. 1974). Reciprocal crosses with all 
taxa, except C. compacta, were performed in the same manner. Twenty crosses 
were performed between C. attem-iata and C. compacta, 9 between C. attenuata 
and C. cuspidata, 16 between C. attenuata and C indecora var. indecora, and 
25 between C. attenuata and C. indecora var. longisepala. 

On the basis of univariate and multivariate analyses, C. 
morphologically distinctive taxon albeit similar to C. indecora, particularly 
var. longisepala. Means, standard deviations, and minimum and maximum 
values for all characters are given m Table 2, as well as LSD test results 
between all taxa for those characters tested. As revealed by the univariate 
analysis, the means of four characters are significantly different (p<0.01) 
between C, attenuata and each of the other taxa (Table 2, Fig. 1). The mean 
of C, attenuata differs significantly (p<0.05) from that of C compacta, C. 
cmpidata, and C. indecora var. indecora, for over half of the characters (15, 16, 
and 15, respectively); and from C. indecora var. longisepala for one-third of 
the characters. 

Multivariate analyses revealed that C. attenuata is distinct from C. compacta 
and C. cmpidata but morphologically similar to C. indecora. In the UPGMA 
analysis, C. compacta and C. cuspidata each formed distinct groupings. How- 
ever, C. attenuata, C. indecora var. indecora and C. indecora var. longisepala did 
not form distinct groupings, but rather one large cluster. 

In the principal component analysis, the first three components explained 
56.4% of the variation (Fig. 2). The remaining variation was accounted for 
by the other factors in 1-4% increments. The first principal component, 
which accounted for 31-1% of the variation, was weighted for characters 
28, 29, 34, and 41. The second principal component, which accounted for 
18.2% ofthevariation, was weighted for characters l4, 21, 31, 32, 39, and 
40. The third principal component, which accounted for 7.1% of the varia- 
tion, was weighted for characters 5, 7, 9, 10, and 37. As in the UPGMA 
analysis, C. compacta and C. cuspidata formed distinct clusters and the three 
remaining taxa did not. The third principal component was weighted for 
characters which dealt with variation in the calyces. For those characters, 
C. indecora var. longisepala was always intermediate between C. indecora var. 
indecora and C. attenuata. 

attmmta differ significantly 
longisepala), and apply to the 

The discriminant analysis yielded probabilities of >.93 that each speci- 
men was appropriately designated, and thus demonstrated that C. attenuata 
and the two varieties of C. tndecora, can be distinguished. 

Interspecific hybridizations. — None of the crosses between C. attenuata and 
any of the other taxa, including both varieties of C indecora, produced fruits 
or seeds. In a related study, populations of C. attenuata were shown to be 
interfertile using these same methods (Prather & Tyrl 1993), and in some 
cases even the same individuals. These intraspecific crosses produced 81- 
92% fruit set and 38-45% seed set. The lack of fruit and seed set from all 
interspecific crosses, but high rate of success of intraspecific crosses, sug- 
gests that C. attenuata is reproductively isolated from the other taxa. 

Taxonomk implications. — Although once thought to be related to C. 
compacta and C. cuspidata, C. attenuata is definitely distinct from these spe- 
cies and should be positioned in subsect. Indecorae on the basis of its fused 
sepals rather than in subsect. Lepidanche as proposed by Waterfall (1971). 
On the basis of the systematic data generated in this study, it is concluded 
that C. attenuata is a distinct species albeit morphologically similar to C. 
indecora. In the absence of reproductive isolation we might treat C. attenuata 
as a variety of C. indecora. 

The addition of C attenuata to subsect. Indecorae brings its total number 
of species to five: C. attenuata parasitizing Iva annua in Kansas, Oklahoma, 
and Texas; C. coryli Engelm. parasitizing a wide variety of hosts in the 
central and eastern U.S.; C. stenolepts Engelm. parasitizmg unknown hosts 
in Ecuador; C. warneri Yunck. parasitizing Fhyla cuneifolia in Utah and 
Arizona; and C. indecora with two recognized varieties: var. tndecora occur- 
ring on a wide variety of herbaceous hosts and widespread in North and 
South America and the West Indies, and var. longmpala parasitizing a wide 
variety of herbaceous hosts in the southwestern U.S., Mexico, and South 
America (Yuncker 1965). Interestingly, C. warnert is also listed as a Cat- 
egory 2 species and is thought to be extinct (1993 PR 58:51159). 

The a 

ithors thank P. Buck, the late J.K. McPherson, and two anony- 

lewers for their helpful comments. Appreciation is also extened to 

the cura 

ors who kindly loaned specimens oi Cuscuta for study. This re- 

search w 

is supported by the U.S. Fish & Wildlife Service (Project Number 



Abacus Concepts. 1992. Statview. Abacus Concepts, Inc., Berkeley, CA. 

Beliz, T.D.C. 1988. A revision oiCmmta sect. Clastogrammica using phenetic and cladisti 
analyses with a comparison of reproductive mechanisms and host preferences in specie 
from California, Mexico, and Central America. University Microfilms International 

'iNKAVA, D.J., R.K. Brown, J. H. Lindsay, and L.A. McGill. 1974. K 

number report XLIV. Taxon 23:373-380. 
'RATHER, L.A. 1990. The biology oiCuKuta attenuata Waterfall. Unpub 

Okla. St. Univ., Stillwater, OK. 
'rather, L.A. and R.J. Tyrl. 1993- The biology oiCumita attenuata Water 

Proc. Okla. . 

• ocular pla, 

Radford, A.E., W.C. Dickison, J.R. Massey, a 

nd C.R. Bell 

atics. Harper & Row, NY. 

SAS Institute Inc. 1985. SAS user's guide: St. 

Taylor, R.J. and C.E. Taylor. 1980. C//u///./ 

tttouuit.i St 

Department of Interior, Fish & Wildlife Se 

\Re, Alhuqi 

Tyrl, R.J., J.L. Gentry, P.G. Rlsslr, and J.) 

C lUKkill. 

evaluation on proposed and candidate end. 

igered plani 

report to the U.S. Department of Interior, I 

Watereall, U.T. 1 97 1 . New species oiCusana , 

nd Phlox fron 


Yuncker, TG, 1920. Revision of the North A 

-nerican and 

Illinois Biol. Monogr. 6: 1-141. 

Yuncker, TG. 1932. The genus CuKuta. Bull 

Torrey Bot. 

Yuncker, TG. 1965. Cuscuta. North America 

1 Flora. Ser. 

Institute Inc. Cat 
port. Submitted i 

, NM, 



. NESOM and 

Depart we 


;;/ of Botany 
■ty of Texas 


PI El 


Amttn, TX 78715, U.S. 


A newly di 

scovered po 

pulation of Dmv?: 

from the Siern 

1 Madre 



vlexico, IS de^ 

illustrated here as 

D. mexkana 

sp. nov 


xpands the ^ 

liree species 

and It Tfii 

■St of the genus k 

nown to occui 

: in Me 


The new s] 

.imilar to D. 

pa/»sim of 

the eastern Unite 

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ncted D. ,.a 

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from D. na 

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ra, reci 


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e iconografiE 


:omo D. mexk. 

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ite en la 

IS ran 

las jove 

nes y ei 

a el enves'de las 

ocadentalis i 


I tubo 

mas largo y mn 



;ortos, y esti 

lo y fib 


tos mas 


. Los dos sisten- 


:)s pot 


iO kilometre; 

s. Dtna 


na esta alejada , 

The genus Dma L. has previously included only two species, D. palustris 
L., scattered but widespread in the eastern United States and adjacent 
Canada, and D. occidentals A. Gray, endemic to six counties of the San 
Francisco Bay region in west-central California (McMinn & Forderhase 1935; 
Vogelman 1953; NevUng 1962; various floristic manuals; Fig. 1). On a 

botanical reconnaissance in September 1994 in the Sierra Madre Oriental 
of Tamaulipas, Mexico, we observed a population of low shrubs that we 
recognized as Dtrca, but the plants were sterile and not identifiable to spe- 
cies. In early March 1995 we were able to study the population at peak 
flowering and beginning of fruit. The Mexican plants most closely resemble 
D. occidentalis, but there are several consistent morphological differences 
between them and they are geographically disjunct by more than 2500 
kilometers (California/Tamaulipas, see Fig. 1). The closest known popula- 
tions o{ Dirca palustns, at the southwestern corner of its range in eastern 
Louisiana, Arkansas, and southeastern Oklahoma, are more than 1100 ki- 
lometers disjunct from the Mexican plants. We believe that it is justifiable 
and desirable to recognize this Mexican population as a species distinct 
from both of its more northern relatives. 

exicana Nesom & Mayfield, sp. 

. (Fig. 2) 

Woody shrubs 6-20 dm tall, averaging ca. 16 dm, from a single t 
branched near the base, with spreading, flexuous branches, the stem 
3(-5) cm wide near the bcise of the plant, the ultimate branches 2-3 
wide, bark smooth and grayish to reddish-brown, the current year's gn 

gle flower, opened 

462 SiDA 16(3) 1995 

persistently loosely and sparsely pubescent, glabrous below. Leaves 
deciduous, alternate, entire, at maturity broadly elliptic to slightly ovate, 
basally rounded, 4-8 cm long, 2.5-6.0 mm wide, 1.3-1.7 times longer 
than wide, the largest distally situated on the branchlets, glabrous above, 
the lower surface persistently sparsely strigose-sericeous on the lamina and 
along the veins, petioles 1-2 mm long. Buds covered by the enlarged peti- 
ole base, mixed (Rowers and leaves), the apical apparently falsely terminal, 
the 4 bud scales whitish-sericeous, forming a foliaceous, deciduous involu- 
cre to the dowers. Flowers appearing before or concurrently with the leaves, 
sessile in axillary and apical fascicles, deflexed to somewhat more nodding 
at full anthesis, the axillary fascicles invariably producing 3 flowers, the 
apical fascicles sometimes apparently "twinned" with (5-)6 flowers; corolla 
absent, not represented by infracalycular structures; calyx (7-)8-10 mm 
long, petaloid, the tube narrow but slightly widening distally, 5-7 mm 
long, abruptly expanded into a flaring limb 2-4 mm long, the tube/limb 
ratio (of length) (1.5-)1. 7-3.0, the 4 calyx lobes 1.5-3.0 mm long, the 
lobes and limb yellow, the tube lighter yellow and drying creamy; stamens 
8, filaments inserted within the calyx tube essentially at a single level (at 
the throat, 2.0-3.5 mm below the lobe apex), the longest exserted 2-3 mm 
above the calyx lobes, those inserted below the lobes 1-2 mm longer than 
those inserted below the sinuses, the thecae 0.4-0.8 mm long, basifixed; 
"hypogynous disc" or "disc" (sensu Heinig 195 1) a ring of connate, irregu- 
lar, slightly fleshy scales ca. 0.2-0.4 mm high and wide, basally adnate to 
the inner surface of the calyx base; stigma minutely capitate, above the 
level of the pre-dehiscent anthers on a fully elongated style, equalled or 
slightly surpassed by the dehiscing anthers. Fruits drupaceous, 1 -seeded, 
pyriform to ovoid, sessile, green when young, mature fruits not observed. 

Typf,: MEXICO. Tamaulipas, Municipio. Hidal^i;(K al.,n,^ mountainous road from Sra. 
Engracia (Tamaulipas) to Dnlces Nombres (Nuevo LL'on)/Arro\o Ohscuro, 2.0 road mi 
NE of Los Caballos toward Canada El Mimbre, 15.0 road mi w'of lowermost crossing of 
arroyo El Mimbre, 23"59'()9"N, 99"28'37"W, ca. 1800 m, S Mar 1995, G/iy Nesoiii 7863 
with Mark Mayfieldand Greg Anderson (hoi.otyfi;: MEXU; isotvpfs: AAU, ANSM, ARIZ, 

jllection exammed: MEXICO. Tamaulipas: type locality, [sterile, le 
m, MayfieU/2()85 with Nesom (TEX). 

of the locality. — The site at which Dina mexu 
I (Arroyo Obscuro) is a steeply sloping, no 
ding a rocky watercourse. The area is limes 
^he elevation near the road is ca. 1800 met 

zana was obser 
oh-facing, m 
tone with kar 
ers, but plant 

Nesom and Mayfield, a new species of Dirca from Mexico 463 

Dirca extend to at least ca. 20 meters (elevation) above the road and to at 
least 80 meters below it. They may well occur over a broader area than we 
investigated, particularly downslope, but this is the only locality where 
Dirca was encountered along the 35 kilometer road from Sta. Engracia to 
Dulces Nombres. Indeed, we found no other site along this road with a 
similar physical and floristic definition. 

The dominant canopy trees at the site are Carya ovata (P. Miller) K. 
Koch, Pmuspatula Schlecht. & Cham., Pseudotsuga menziesii (Mirb.) Franco, 
and Quercus launna H. & B. Large trees of these reach about 20-25 meters 
in height, with tall Pseudotsuga perhaps reaching 30 meters. Judging from 
stumps in the area, particularly large individuals (presumably o^ Pseudotsuga 
and Pmus) have been removed by logging. Large trees oiCarpinus caroliniana 
Walt, and Uqmdambar styraciflua L. are scattered at the locality but do not 
reach the full height of the canopy. The woody understory includes the 
following: Cornus urbimana Rose, Croton virletianus Muell.-Arg., Garrya 
macrophylla Benth., Ilex aff. rubra S. Wats., Litsea pringki Bartlett, Persea 
pododaema Blake, Philadelphus caldcolus Hu, and Taxus globosa Schlecht. Most 
of the canopy species are deciduous and were just beginning to break bud 
at this site; the understory includes a predominance of evergreen species. 
Arboreal bromeliads, including three species of Tillandsia, are conspicu- 
ous. Ferns form a major part of the herbaceous flora at this site; among the 
most common are species of Adiantum, Woodivardia, Botrychium, 
Phanerophlebta, Polypodium, and Polystkhum. Other common herbs include 
species o^ Chmaphila, Chiropetalum, Goodyera, Sisyrinchium, and Stachys. 

Characteristics of the population and biological observations. — We observed 
800-1000 plants (by estimate) at the Dirca site, where they occur for about 
300 meters along the road and adjacent slopes on the east side of the water- 
course. They tend to be densely clustered in more open-canopy microsites, 

singly and more scattered over a larger area. We did not unearth any plants, 
but there was no indication that they reproduce clonally. A report of rhi- 
zome production in D, occidentalis (McMinn & Forderhase 1935) has not 
been corroborated by more recent observations (Spongberg pers. comm.). 
The entire population of Dirca mexicana is strongly synchronous in 
flowering. Had we arrived 3 or 4 days later, it is likely that we would have 
been unable to observe floral features, as the flowers appear to wither quickly 
with the onset of fruit maturation. The flowers appear to be weakly 
protogynous and there is evidence that they also may be self-compatible. 
Initially, before full development of the calyx, the style (with apical stigma) 
usually is elongated past the level of the anthers. At this time, the stigma 
appears slightly moist and presumably receptive; we also observed that 

many at this stage appear to have pollen attached. At full anthesis, the 
filaments have raised the open anthers to the level of the stigma or slightly 
beyond it. Further, there is little time separating intial receptivity of the 
stigma and dehiscence of the anthers, and little distance between the stigma 
and open anthers. 

Visiting the flowers were one species of bee (individuals ca. 7 mm in 
length) and at least four species of butterflies. Pollination is effective, judg- 
ing from the apparently ubiquitous deposition of pollen on the stigmata of 
pre-staminate flowers. Bees and butterflies were probing inside the flowers, 
and although it seems highly likely that nectar was available to them (prob- 
ably produced by the hypogynous disc), its production was not evident in 
numerous flowers that we examined. Nor could we detect any fragrance 
(D. ocddentalis was noted by Howell [1970} to be fragrant). Fruit matura- 
tion was beginning only on a few plants and we were unable to make an 
estimate of the success of fruit and seed production. 

We did not determine what feature or features account for the remark- 
able rarity o^ Dirca mexicana. All of the other species yet identified from 
Arroyo Obscuro are found in other sites in the same general area — none 
except the Dina could be considered rare (but see comments below regard- 
ing Viburnum). Rarity, however, also is a feature of both other species of the 
genus, particularly D. occtdentalts (Stebbins 1942; Johnson 1994). D'ma 
palustns is widespread but of uncommon occurrence. 

Morphological amipansons. — Dirca mexicana resembles D. ocddentalis in most 
of the features that have been used to distinguish the latter from D. palustris 
(Vogelmann 1953; see key below): vestiture, presence or absence of a pe- 
duncle and pedicel, and the distal morphology of the calyx. The flowers of 
D. mexicana, however, are more similar in general configuration to those of 
D. palustris: both have a relatively long and narrow tube with a shorter, 
abruptly widening limb, the staminal filaments are inserted above the 
middle of the calyx, and the style and anthers are exserted for a relatively 
short length. In D. occidentalism the tube is shorter than the broadly funnel- 
form limb, the staminal filaments are inserted below the middle of the 
calyx, and the style and stamens are long-exserted. Vogelman (1953, p. 80) 
emphasized the taxonomic usefulness of the level of filament insertion, which 
in turn is indicative of the throat position and flower shape: "In most in- 
stances this character alone is sufficient to distinguish the two species." 
The comparative illustrations furnished by Vogelmann, however, do not 
accurately represent this difference, nor does the detailed illustration of D. 
palustris in Cronquist (1981, p. 635); those by Holm (192 1) are more simi- 
lar to our own observations. Differences between the new species and D, 

Bud scales with bro 

wn or reddish-br 

own pi 

ibescence; young twigs ai 

nd both 

leaf surfaces complf 

itely glabrous; fl. 

3wers I 

ind fruirs pedicellate, th. 

e whole 

cluster often pedun 

.culate; calyx ma 

rgin merely crenulate-undulate 

;, with- 

out distinct lobes;. 

=astern United St 




Bud scales with wh 

■itish pubescence 

; youn, 

g twigs and abaxial leaf: 


persistently pubesci 

;nt; flowers and f 

ruits se 

ssile; calyx distinctly anc 


Bay region of Califori 

iia or northeastern Mexic 

2. Calyx tube 2-4 r 

nm long, broade 

ned inl 

:o a broadly funnelform 1 

imb 4- 

6 mm long; stan 

amal filaments u 


below the middle of the calyx. 

3_4(-5) mm at 

)ove th 

.e flower at maturity; st 

yle and 

stigma (l-)2-4 

mm above the a 


at maturity; San Franci 

SCO Bay 

Bwgeographk pattern. — It does not seem possible at this point to provide 
a morphologically based hypothesis of relationship among the three spe- 
cies ofDirca, because the genus appears to be relatively isolated, its closest 
relatives (and thus the evolutionary polarity of character states) difficult to 
specify (see Domke 1934; Nevhng 1959). Still, it is surprising to find a 
greater overall similarity between D. mexicana and D. occidentalism in view 
of the well-known pattern of close relationship and disjunction between 
species of the eastern and southeastern United States and the sierra of north- 
eastern Mexico (Miranda & Sharp 1950; Graham 1973). In the immediate 
area of Dirca mexicana are numerous species that are disjunct from their 
primary range in the eastern United States (e.g.): Carpinus caroliniana, Carya 
ovata, Chimaphila umbellata (L.) W. Barton, Desmodium glutinosum (Muhl. ex 
Willd.) Wood, Liquidambar styraciflua, Pedicularis canadensis L., and 
Polystichum acrostichoides (Michx.) Schott. In addition, there is a rare and yet 
undescribed species oi Viburnum (Nesom in prep.) in the close vicinity that 
apparently is most closely related to V obovatum Walt, of the southeastern 
United States. Another recently recognized, disjunct species from the same 
area of Tamaulipas, Scrophularia sp. nov. (May field and Nesom submitted), 
is closely similar to S. marilandica L. of the eastern U.S. species and S. 
californica Cham. & Schlect. of the Pacific region. Taxus globosa, one of the 
understory species at the Dirca site, also has close relatives widely separated 
in the eastern and western United States. 

Biotic disjunctions between the eastern and western United States, similar 
to that in Dirca, are well known though not particularly common (e.g., 
Sharp 1951; Wood 1970), and other examples can easily be added (e.g., 
Sericocarpus, lonactis). This pattern is emphasized by the recent discovery of 

466 SiDA 16(3) 1995 

a second species ofNeu^isia in California (Shevock et al. 1992), both species 
extreniely rare. In contrast, clearly established disjunctions of extant plants 
from the western United States to the Sierra Madre Oriental of Mexico are 
unknown to us. Numerous disjunctions exist between the Sierra Madre 
Oriental and S.M. Occidental, but most or all of these appear to be deriva- 
tives of the general pattern described by McVaugh (1952), where the evo- 
lutionary antecedents occupy a more southern position. A few Mexican 
species have a broken but still somewhat continuous distribution from the 
eastern sierra through northern Coahuila into the mountains of southwest- 
ern Texas and southeastern New Mexico (e.g., Populus tremuloides Michx., 
Pseudotsuga menziesii [Mirb.} Franco, and see Nesom 1993). 

In the broadest view of the origin of Dm-^ mexicana, the simplest hy- 
pothesis is that it belongs to the floristic element with its closest evolu- 
tionary ties to the flora remaining in the southeastern United States, the 
geographic continuity between them probably established during the middle 
to late Miocene (Graham 1973). The disjunction between D. palustns and 
D. ocadmtaln may be considerably older, as a vegetation probably includ- 
ing these species was spread across North America through much of the 
Tertiary, beginning as early as the Eocene (Graham 1972, 1993). In a con- 
trasting view, Axelrod (1975) hypothesized that the closely related dis- 
juncts found in eastern Mexico, the Appalachians, and the West coast of 
the United States represent remnants of a continuous forest earlier spread 
into Mexico and more simultaneously fragmented as a result of a spreading 
dry climate in the mid-Oligocene. Evaluation of the latter theory is diffi- 
cult at present because of the lack of evidence for northern temperate ele- 
ments in Latin American prior to the Late Tertiary. These elements have 

not been rec 

covered from 

the Oligo-Mi 

ocene Sim 


1 Group 

of Chiapas, 

Mexico; Q^ue 

reus first appe 

■ars in Panam; 

a in the M 


ocene, ai 

id in South 

America A//, 

ins arrives by 

about one mi 

Uion years 

ago a 

nd Q^uen. 

us not until 

about 340, ( 

)()() years age 

). Many east( 

;rn North 


rica disj 

uncts were 

present in 

eastern Mc 

_^xico by th< 

; middle 


cene, a 

nd global 


ature history 

suggests that 

cooling in 

the middle M 

iocene may 

have been ar 

1 appropriate 

time for thei: 

r principal 


We thank Greg Anderson for help with collecting, observations on floral 
biology of the new species and his comments on the manuscript. We are 
grateful to Billie Turner, Steve Spongberg, Alan Graham, and Beryl Simpson 
for comments on the manuscript, to John Taylor, Gene Wofford and Bob 
Krai, and Sidney McDaniel for information on Dirca in Oklahoma, Ten- 
nessee, and Mississippi, respectively, to Linda Vorobik for the illustration, 
and to Jaime Hinton for his encouragement and company on fall and spring 


illecting trips and for the example of his 
nuing vigor. 

y Press New York 

1934 Untersucliun^en Liber die s> stem itischeundgeographischeGhederun^ 

1972 Outhnt ot tlu oiuin in 1 historit d reLo^ninon of flonstic iffinitie: 
Asii ind eastern Noirh AmeiRi In A Crihim (cd ) rionstics ind paieo- 

_ 197^ Historv ol the irboitscuu tempLrite element in the northern Latir 
bioti In A Grihim cd Vci^etuion md Ve^et ition History ol Northerr 


\ lU, 199 

1 PliUuolthe\Ve-,tL. 



, ,) M I [ ind 

> Mex 

4LSOM Submitted 

A new 


N 1 1 I u.d 1 

} [OR 

DLRHAS, 19.5 ^ 

Jotes on 


Midiono ■> 


M< Vm 


1 Hut 

sttd phyiot^env 



M 1 'ind A ) 

Rr 1950 Ch^ru 



"(Vj' ' 199'' 


e species ot Astc 

! I A^ste 


null Mexico 


ologia 74 296-3 




' The 

m tlie 

I Dma ouidentalis 

J Dina ouidentalts (Thymeleaceae). 
;n the southern Appalachians and 


Fautin, Daphne Gail, Douglas J. Futliyma, and Frances C. James. 1994. 
Annual Review of Ecology and Systematics, Volume 25. (ISSN 
006-4162, hbk.). Annual Reviews, Inc., 4139 El Camino Way, Palo 
Alto, CA 94303-0139. $47.00 US, $52.00 elsewhere. 686 pp. 

"Chapters in this volume directly relevant to application of ecological and systematic 
knowledge are those by Schulze et al., who apply new technology to some pressing global 
issues, Chambers and MacMahon, who compare natural and managed systems, and Leigh 
Brown and Holmes on HIV. Holt and Lawton and Kuris and Lafferty, among others, ex- 
plicitly relate principles they discuss to human concerns. Linkages between and among 

and systematics, is implicit in many chapters; it is explicit in those by Moran on complex 
life cycles, Moreno on genetic architecture, Raff et al. on metazoan phylogeny, and 
Vnjenhoek on unisexual fishes, and in that by Vermeij entitled "The Evolutionary Interac- 
tion among Species: Selection, Escalation, and Coevolution." "Three chapters concern sys- 

Mesembryanthemaceae by Ihlenfeldt." There are 25 chapters: 1) Algal Nutrient Limita- 
tion and the Nutrition of Aquatic Herbivores; 2) Genetic Architecture, Genetic Behavior, 
and Character Evolution; 3) Molecular Approaches to Population Biology; 4) Unisexual 
Fish: Model Systems for Studying Ecology and Evolution; 5) Cooperation and Conflict in 
the Evolution of Signal Interactions; 6) Evolutionary Biology of Human Immunodefi- 
ciency Virus; 7) Metapopulation Dynamics and Genetics; 8) Community Structure: Larval 
Trematodes in Snail Hosts; 9) The Evolutionary Interaction Among Species; Selection, 
Escalation, and Coevolution; 10) The Systematics of Coral Genus Acropora: Implications of 
New Biological Findings for Species Concepts; 1 1) A Day in the Life of Seed: Movements 
and Fates of Seeds and Their Implications For Natural and Managed Systems; 12) The 
Evolution of Vocalization in Frogs and Toads; 13) Phylogeny of the Legume Family: An 
Approach to Understanding the Origins of Nodulation; 14) Using DNA Sequences to 
Unravel the Cambrian Radiation of the Animal Phyla; 1 5) Genetics and Ecology of Whales 
and Dolphins; 16) Fisheries Ecology in the Context of Ecological and Evolutionary Theory; 
17) Ecology and Evolution of Reproduction in Milkweeds; 18) The Nature and Conse- 

Evolutionary Ecology: Ariolis Lizards as Model Systems; 20) The Ecological Consequences 
of Shared Natural Enemies; 21) Diversification in an Arid World: The 
Mesmembryanthemaceae; 22) Genetic Divergence, Reproductive Isolation, and Marine 
Speciation; 23) Adaptation and Constraint in Complex Life Cycles of Animals; 24) Male 
Parental Behavior in Birds; 25) Relationships Among Maximum Stomatal Conductance, 
Ecosystem Surface Conductance, Carbon Assimilation Rate, and Plant Nitrogen Nutri- 
tion: A Global Ecology Scaling Exercise. A subject index is included. 





Department of Botany 

University of Texas 

Austin, TX 78713. U.S.A. 


3se examination of herbarium specimens coupled with fieW 

/pic by various ; 
observations ir 

,at Hc,plol^hyto„ 

1 Chi< 


aplophytinae. Distribution maps and a key to the species are 


EY Words: Hapbphyton, Alstoma, Haplophytmae, Alstominae 

, Apocynaceae 

De CandoUe (1844) described Haplophyton as a monotypic genus based 
upon H. cimictdum. Benson (1942) added a variety to the species, var. crooksii, 
later elevating it to the rank of species (Benson 1943) when material of 
mature seeds was examined. Benson (1954) regarded H. crooksii as a dis- 
tinct taxon because of its smaller leaves and smaller discontinuously ridged 
seeds. Various authors gohnston 1990; Kartesz 1994; Leeuwenberg 1994; 
Pichon 1950), however, disregarded these differences and recognized only 
H. dnnctdum. My exammation of the specimens housed at LL, TEX coupled 
with field observations in Chiapas, Mexico, however, has uncovered new 
evidence that strengthens the recognition of two distinct species. 

The genus Haplophyton is placed in the subfamily Plumerioideae because 
of its undifferentiated free anthers and mdole alkaloides. These characters, 
in particular the indole alkaloides, are regarded as indicative of a mono- 
phyletic lineage (Leeuwenberg 1994). In addition, sinistrorse aestivation 
of the corolla (overlapping of the petals in bud to the left) distinguishes the 

Plumerioideae from the Apocynoideae, which has dextrorse aestivation (over- 
lapping of the petals in bud to the right) (Leeuwenberg 1994). The 
aestivation oiHaplophyton, however, has been under considerable question. 
De Candolle (1844) described Haplophyton as sinistrorsely contorted, while 
Bentham and Hooker (1873) listed the taxon as having a dextrorse 
aestivation. Pichon (1950) also pointed out that Haplophytou had a "curi- 
ous" dextrorse aestivation. Woodson (1938) on the other hand reported a 
sinistrorse aestivation for Haplophyton, as did Standley and Williams (1969), 
and Correll and Johnston (1970). Benson (1942, 1943, 1954) did not de- 
scribe the aestivation. An examination of herbarium specimens, however, 
reveals that the above reports are all correct. The southwestern United States 
and northern Mexican (Fig. 1 ) members o{ Haplophyton have an aestivation 
to the left, while the Guatemalan and southern Mexican (Fig. 2) members 
have an aestivation to the right. The correlation of this character with the 
allopatric distribution of the two populations provides strong evidence that 
Haplophyton is comprised of two distinct species. My field studies in El 
Chorreadero, Chiapas {Williams & Plum, 95-34) confirm that southern popu- 

When de Cando 

Oaxaca. Topotypes from this regi( 

1844) described Haplophyto? 
" • ^ ' type a spec: 

ikidum he reported a 
n from Tehuantepec, 

Because I have not had an opportunity to examine the type, its aestivation 
remains at present undetermined. I suspect, hov^^ever, that it is to the right 
and that the reports of a sinistrorse aestivation are either from specimens 
examined from the north or are the result of the presupposition that all 
Plumerioideae flowers have aestivation to the left. 

Differences in the corolla tube and calyx dimensions have not until now 
been discussed as segregating factors between the two species, but my ex- 
amination of herbarium specimens shows that H. cimicidum possesses a longer 
corolla tube than H. crooksii, as well as smaller calyx lobes. An examination 
of the seeds, in particular their size and surface structures, also supports 

that seems to tentatively hold true is the color of the coma which is typi- 
cally golden yellow in H. cimiadim and charcoal grey in U. crooksii. 

The populations in Guaymas, southwestern Sonora, superficially resemble 
H. cimicidum in leaf size; accordingly Benson (1943, 1954) recognized them 
as such. A close inspection of these individuals, however, shows that their 
aestivation is to the left and that the seeds are 6-8 mm long, placing them 
with H. crooksii. 

With its sinistrorse aestivation, larger calyx lobes, smaller corolla tube, 
smaller seeds, and isolated geographical range, it seems evident that 

SiDA 16(3) 1995 

Haplophyton crooksii constitutes a distinct element from H. cimkidiim and, 
consequently, necessitates the recognition of two species for Haplophyton. 

mm wide; corolla tube 8-10 mm long; sepals 3-5 mm long, 
the length of corolla tube; seeds 8-1 1 mm long, vertical ri 

. Petals of the bud overlappmg to the left; leaves 1 1-35 (50) m 
mm wide; corolla tube 6-8 mm long; sepals 4-1 1 mm long, I 
equal to half the length of corolla tube; seeds 6-8 mm long, \ 

western United States and northern Mexico 

(TEX) El ( 
34 iTL\) i 

/ (TEX) ^2 km S of 
/ (TLX) 01 en irc is 9- 
J59 Knt I^l (TLX) 

2. Haplophxt<Mi crooksu (J D Benson) L D Benson Amcr J B^ 


I EX) T.x\s El Piso Co Eiinkhn Mts 1 1 mi V( jct Trins 
'» and Gitewav S (M >3 N 106 2^ W) southern exposuie of 
-x 26 Aug 19-^8 WorthiN^t,,? }2(raLX) Hudspeth Co hme 

Williams, Notes on Haplophyton 47 3 

basin in the Quitman Mts, 17 Apr 1976, 

mouth of narrow canyon draining S slope of the W Chinatis, among igneous boulders, 

4800 ft, 14 Jun 1977, Butterwick 5857b (TEX). 

MEXICO. Chihuahua: ca 20 km ENE of Ciudad Jimenez, in limestone arroyo in can- 
yon NW of summit of Sierra de Chupaderos (27°12'N; 104°43'W), 5100 ft, 2 Oct 1973, 
Henrickson 13765 (TEX); ca 31 mi NW of Julimes in SW facing canyon above Rancho El 
Recuerdo in Sierra de Carrasco (28°47'N; 105°09'W), 4400 ft, 15 Sep 1973, Henrickson 
12941 (TEX); 46 km W of Ojinaga on hwy to Chihuahua City and S 2 km to the deep 
canyon of Rio Conchos in the Sierra de Pegiiis (29°32'N; 104°48'W), 1000 m, 20 Oct 

1972, Chiang ei al. 9757 (TEX); S slope and top of Sierra del Roque, NNE of Julimes 
approached from Mina Las Playas via Rancho El Saucito (28°39'N; 105°18'W), 19 Jun 

1 973, M.C.Johnston et al. 11386B (TEX). Coahuila: S side of Cerro San Jose de las Piedras 
(28°42'N;102°51'W), 1000-1400 m, 8 Jun 1972, C^/^«^e^^/, 7355 (TEX); 0.5 km E of 
MatrimonioViejo,E end of limestone ridge (27°08'N; 103°07'W), 1125 m, 2 Sep 1972, 
Chiang 9122 (TEX). Sonora: Sierra Bojihuacame SE of Obregon, 800-2000 ft, 17-25 
Oct 1954, Gentry 14482 (TEX); talus slope of low basaltic hill, 15 mi S of La Palma, 
between La Palma and Guaymas, 2 Sep 1 94 1 , Wiggins 226 (TEX); Rio Mayo area, hill with 
microwave tower, 9 mi NW of Alamos and 4 mi SSW of Piedras Verde (27°07'N; 
109°02'W), 500-700 m, 6 Sep 1989, Sanders 9435 (TEX); Canon de Nacapules, 6 km NE 
of Bahia San Carlos, deep riparian canyon, 19 Oct 1984, Felger 84-122 (TEX). 

In his classification oi Haplophyton, Pichon (1950) erected the monogeneric 
subtribe, Haplophytinae. He discussed its affiliation with the 
Catharanthinae, but ended by questioning the relationship, remarking that 
"the lignification of the branches [in Haplophyton'] seems to be rather rapid 
and the plant has nothing in common in this point of view with the 
Catharanthinae" (Pichon 1950; p. 1 6 1). Leeuwenberg (1994) subsequently 
positioned Haplophyton with Alstoma R. Br. in the Alstoniinae, near the 
Catharanthinae, on the basis that both genera are the only two taxa in the 
Plumerioideae to possess hairy seeds. Seed pubescence in the Alstoniinae, 
however, is heteromorphic. Haplophyton has seeds with both apical and basal 
coma, while Alstonia has seeds with either membranous ciliations (sect. 
Tonduzia) or with both apical and basal coma (sect. Monuraspermum) 
(Monachino 1949). 

The dextrorse aestivation found in Haplophyton, suggests a close rela- 
tionship with Alstonia, which also displays dextrorse aestivation. Indeed, 
Haplophyton cimkidum may be related to the Oceanic species of Alstonia 
(sect. Monuraspermum), which have flowers with dextrorse aestivation and 
seeds with both apical and basal coma (Monachino 1949). Presently, how- 
ever, it is not known whether or not dextrorse aestivation in Haplophyton is 
ancestral or derived. 

Alstonia is the only other genus in the tribe Plumerieae, besides the gen- 
era of the Catharanthinae (excluding \tnca (Lawrence 1959)), to retain a 
reflexed membranous appendage basal to the stigma throughout its ontog- 
eny (Pichon 1950). Woodson (1928) noted, however, that a swollen region 

474 SiDA 16(3) 1995 

occurs below the stigmas oi Haplophyton. In his view, this character prob- 
ably represents a primitive stage of the more complex appendage found in 
the Catharanthinae. Preliminary studies of Haplophyton herbarium speci- 
mens, by the author, reveal that the style heads of immature buds have a 
membranous skirt below them. This feature, however, was not present in 
mature (opened) flowers in either the field or herbarium specimens. 

Pichon (1950) also suggested an affiliation between Haplophyton and 
Anechites, stating that the two genera are unique in the family by having 
hairs with multicellular bases (Fallen 1983). Pichon, however, placed each 
of these genera in monotypic subtribes, suggesting that their relationships 
were unresolved. Anechites was subsequently positioned in the subtribe 
Condylocarpinae of the subfamily Plumerioideae (Leeuwenberg 1994). 

Haplophyton differs from Alstonia sect. Monuraspermum in being a 
suffruticose herb (vs. tree), having alternate (vs. whorled) leaves, hairs with 
multicellular bases (vs. simple), no nectary (vs. an annular nectary), soli- 
tary flowers (vs. cymous), and its restriction to the New World (vs. Oceania). 
These differences suggest that Haplophyton is not related to Ahtonia. Until 
further evidence is presented, the most suitable and conservative course 
favors placing Haplophyton in the monogeneric subtribe Haplophytinae 
(Pichon 1950) of the tribe Plumerieae, and not with Ahtoma in the 
Alstoniinae (Leeuwenberg 1994) whose resemblance to Haplophyton is at 
present questionable. 

I am grateful to Carol Todzia, James Henrickson, and the two anony- 
mous reviewers for their attentive editing of the manuscript, and to my 
companion Jennifer Forrest for translating Pichon's work from the original 
French. B.L. Turner provided helpful comments. Cooperation from F, and 
TEX-LL was also much appreciated. Finally, I acknowledge my friend Jon 
Plum, for having the tenacity to accompany me to Chiapas, despite the 




L.D. 1 

942. Notes on 

the flora 


;ona. To: 

Bot. 3 


^^ . 

ndR.A.DARROw. 1954 



icy of Arizona,' 



I. Pp. 2i 

Bentham, G. ai 


<. 1873. 


laceae. 1 


.London. 2:722-723. 


E, A. E 


laceae. Ii 

n: A. P. c 

le Cand( 



^iHs. Forc.n, M: 

asson & 

cie, Par 

IS. 8:41. 

Williams, Notes on Haplophyton 

CoRRELL, D.S. and M.C. Johnston. 1970. Manual of the vascu 

Research Foundation, Rennet, Texas. R 1211. 
Fallen, M.E. 1983. A systematic revision oi Anechttes (Apocyn 

Johnston, M.C. 1990. The vascular plants of Texas; A list up- 
Canada, and Greenland, 2nd ed. Timber Press, Portland, Ore 

AWRENCE, G.H.M. 1959. Vm-^ and 

1 Catharanth, 

w. Baileya. 7:11.^ 

EEUWCNBERG,A.J.M. 1994.Taxaol 

r the Apocyr 

,aceae above the ^ 

Wageningen Papers 94(3):45-60 

iCHON, M. 1950. Classification des . 


: XXVIII, Supple 

Mem. Mus. Natl. Hist. Nat., Ser 

. B.,Bot. 1:: 


osATTi,T 1989. The genera of suh 

>order Apocy 

-nineae (Apocyna 

in the southeastern United States 

;, Apocynace 

•ae. J. Arnold Arl 

c Botanical Garden, New Yoi 


Rice, Elroy L. 1995. Biological Control of Weeds and Plant Diseases. 
Advances in Applied Allelopathy (ISBN 0-8061-2698-1, hbk ) 
University of Oklahoma Press 1005 Asp Avenue, Norman, OK 73019, 
(405)325-511] $55 00 4^8 pp, ] 4 figures 

Bioloj^ualConliol oj W c A /;/ / P/ „t D, /, \ h //;,m ni Applud Mldop ,thy is a timely 
study, by I world renowned uitlu)nc\ of clu piodLRtion b> phnts ind micioorgamsms, 
of compounds that ifkct clu ^lowrh ht ilth md popuhtion biolo,) of other plants and 
microorgamsms Ulo^ L focuses on neu dcxclopmcnts in tUelopuhy in agriculture 

the last decade, Ric 

;)t Phnts 6) Allelopith) in the Biologic il Control 
) Allelopathy in the Biological Control of Plant 

Frey, Kenneth J., Ed. 1995. Historical Perspectives in Plant Science. 

(ISBN 8138 2284 X, hbk ) Iowa State University Press 2121 S 
State Avenue Ames, lA 50014 8300 1 800 862-6657 $44 95 216 

md Gerty Con Wird 7) Phnt Breeding— A Vitil P 
:^)Lulitj and Production Efhciency b> John W DudI 
Phnt Breeding Methodoloi,y byNcilF Jensen 



Forest Plant Laboratory 

Southwest Forestry College 

Kunming, Yunnan 650224, CHINA 


'longensts G.S. Fan & L 
Kmsleyanus Schneid. v 

,L. Deng sp.nov. from Yur 


and Rhamnus hemsleyanus Schneid. v^t paucinervatus G.S. Fan & L.L. Dent; var. nov. from 
Wuyi Shan Mountain, Fujian, China, are described and illustrated. 
Key Words; New taxa; Rhamnaceae; China 

Se describen e ilustran Sageretiaynnlongemis G.S. Fan et L.L. Deng del condado de Yunlong 
en Yunnan (China) y Rhamnus hemsleyanus Schneid. var. pauanervatus G.S. Fan et L.L. Deng 
var. nova de la montafia Wuyi Shan en Fujian (China). 

Sageretia yunlongensis G.S. Fan & L.L. Deng, sp. nov. (Fig.l) 

foliis oblongo-ellipticis, 10-15 cm longis, 4.5-6.5 cm latis; axibus inflorescentiarum et 

Shrub 4 m tall, unarmed; branchlet cylindric. Leaf thick and papery, 
oblong-elliptic, 10-15 cm long, 4.5-6.5 cm broad, obtuse-rounded to short- 
acute at apex, rounded at base, pubescent beneath; the midrib pubescent 
above; the lateral veins 7-8 per side, impressed above, raised beneath; peti- 
ole thick, 14-20 mm long, densely pubescent. Inflorescence paniculate, 9- 
10 cm long in fruit; axis of inflorescence densely pubescent, persistent in 
fruit; flower unknown. Fruit globose, 3-4 mm diam.; pyrenes 2-3; stem of 
fruit 2-3 mm long. 

Typi;: CHINA. Yunnan: Yunlong Xian, 1300 m alt., Yin Wu-Yuan et al. 161, 26 Oct 
1987 (HOLOTYPc: SWFC). Paratypes: CHINA. Tibet, Motuo Xian, Tibet Team 74-1702. 17 
Aug 1974 (KUN). 

This new species is similar to Sageretia henryi Dumm. & Sprag., but it is 
easily recognized by its oblong-elliptic, 10—15 cm long, 4.5—6.5 cm broad 
leaf, and axis of the inflorescence, the lower leaf surface, the midrib above, 
and the petiole with pubescence. 

Rhamnus hemsleyanus Schneid var paucinervatus G.S. Fan & L.L. Deng, 

imilcir CO vai hem\h')cini 
> (4— '5 pairs) and glabic 





Department of Biological Sciences 

Northern Kentucky University 

Highland Heights, KY 41099-0400, U.S.A. 

ar chronology of interaction bet 
; of arboreta, botanical gardens, 
;ntual naturalization. The specie 

introduccion y eventual naturalizacion de esta planta. Se describe la biologia de esta especie, 

In an effort to understand how non-indigenous plants occupy new geo- 
graphic areas, two processes have received the most attention: population 
spread as documented from herbarium records (Forcella 1985) and popula- 

(Mack 1985). Although most plant mvasions result from accidental or in- 
to human cultures and the historical and extant exchange systems contrib- 
uting to plant invasion. A description of these systems, when they operate 
with plants that eventually escape and become naturalized, may be useful 
for understanding rates and areal extent of the invasion process. 

We present here a 150-year chronology of events that eventually led to 
introduction and naturalization of the eastern Asiatic shrub Lomcera maackii 
(Rupr.) Herder (Caprifoliaceae), Amur honeysuckle, in North America. 
Throughout most of this time L. maackii was highly valued in gardens and 
conservation plantings. However, the tendency of the species to naturalize 
and spread beyond points of original introduction established it as a woody 
"weed" of concern in eastern U.S. The documented history of interaction 
between Amur honeysuckle and people is both extensive and varied. The 
chronology given here may be of value as regulatory decisions are made 
about future plant introductions. 

4S0 Sum 16(3) 1995 

The addition of L. mctcickii to the alien flora of North America can be 
traced to three historical interactions between the plant and Homo sapiens: 
(1) Discover the plant, classify it, and describe it as a member of the flora of 
eastern Asia; (2) Introduce the plant to western horticulture for its attrac- 
tive foliage, abundant flowers, and showy fruits; and (3) Use the plant to 
achieve conservation goals, e.g., soil stabilization and/or wildlife-habitat 
improvement. These three interactions contributed to widespread intro- 
duction, thus necessitating a fourth and final one: control or remove the 
plant from the many biotic communities that have been invaded. 

From the Pacific to Western Europe: The Russian Role 

In the mid 1 9th century, Russian possessions in coastal eastern Asia were 
all north of the 55th parallel, a somewhat less than hospitable region. To 
extend their holdings into more favorable areas, the Russians initiated a 
series of southward explorations into the relatively greener pastures of 
Manchuria (Bretschneider 1898). This area, then "but loosely held in the 
feeble grasp of the Chinese government," had excellent harbors and abun- 
dant resources, including timber. Among the targets was the territory north 
of the Amur River and that bounded by the Amur and Ussuri rivers, the 
Sea of Japan, and the Korean frontier. Russia eventually annexed these lands 
by treaty from China in 1858 and I860, thus extending its domain south 
to the latitude of present-day Vladivostok. 

One of the scientific expeditions sent to explore the valley of the Amur 
began at Irkutsk in April 1855, returning to Irkutsk 9 months later 
(Bretschneider 1898). Accompanying the expedition was a naturalist, Ri- 
chard Maack (1825-1886), professor in the Gymnasium of Irkutsk. Maack is 
remembered today primarily in the name of a genus of Fabaceae, Maackta, 
and in the specific epithets of several species, including one in Lonkera. 

Among the species that Maack found along the Amur in June was the 
yet-to-be-described Amur honeysuckle; he made but a single collection of 
the plant — in the Bureja Range north of the Amur about midway between 
Khabarovsk and Blagoveshchensk (Maximowwicz 1878; Ruprecht 1857). 
His specimens of woody plants from the trip were sent to St. Petersburg, 
where they provided the basis for part of the first publication devoted to 
plants of "Amurland" (Ruprecht 1857), that area on both sides of the Amur 
between ca. 42° and 55° north and 131° and 141° east (Maximowicz 1859). 
In that work the honeysuckle was described as a new species, Xylosteum 
maackii, by Ruprecht (Ruprecht 1857). Maack (1859) gave an account of 
his journey along the Amur; in this work is the first published illustration 
(at least in western literature) of Amur honeysuckle (Fig. 1). The species 
was soon included in Maximowicz's Pnmitiae florae Amurensis (Maximowicz 

1859), the first flora of Amurland, written after Maximowicz's first trip 
(1853-1857) to eastern Asia. Five years after publication of the flora, the 
species was transferred to Lonicera as L. maackti (Ruprecht) Herder (Herder 

Maximowicz did not see the plant in nature before he wrote Pnmitiae 
florae Amurensis. However, in a later expedition (1859-1864) he obtained 
specimens of it for St. Petersburg from five localities (Herder 1878): near 
the mouth of the Amur, i t St. Olga Bay, and at three sites near Vladivostok. 

We do not accept repc^rts (e.g., Bretschneider 1898) that Maack "intro- 
duced" L. maackii to culti ration at the St. Petersburg Botanic Garden; Boom 
(1959), Rehder (1949b), ind Wyman (1969) even date this event as "I860." 
We conclude that Maack did not bring seeds or living plants of L. maackii 
to the Garden from eithe • of his trips (1855, 1859). If he had brought back 
such propagules, these would have produced blooming plants much earlier 
than 1883, the date recorded in Gartenflora (Regel 1884) for the first Euro- 
pean flowering of the plait, at the St. Petersburg Garden. This beginning 
of the plant's ascent in western horticulture was some 24 years after Maack s 
last return from eastern / sia. The species comes into flower in 3 to 5 — not 
24— years from seed (Lo "enz et al. 1989). The flowering of other eastern 
Asiatic woody plants raised from seeds that Maack did send to St. Peters- 
burg (e.g., species oiClen'atis, Pyrus, Deutzta, and Syringa) was reported in 
the early 1860s (Bretschr eider 1898). 

According to Thatchei (1922), plants of L. maackii were introduced to 
St. Petersburg from Manchuria— introducer unknown— in 1880; these 
plants could well have be;n the ones that came into flower in 1883. (That 
seeds were the introduced p "opagules was maintained by Anonymous {1924}.) 

Regel's (1884) report cf the flowering of L. maackii in St. Petersburg in 
1883 was soon translated abridged, and published in horticultural works 
in England (e.g., Anonyinous 1884a, 1884b; Nicholson 1888) and the 
United States (Bailey 190 3; Davis 1899); the authors of these reports obvi- 
ously had not seen living ^^xamples of the plant. However, within a decade 
after the 1884 article, deiailed morphological data obtainable only from 
//i^^ plants were published in Germany (Dippel 1889; Koehne 1893), indi- 
cating cultivation in that country. In 1896 at least one German nursery 
grew L. maackn (Cole, fers. comm.). The National Botanic Gardens, 
Glasnevin, Dublin, Ireland, purchased plants of Amur honeysuckle from 
the French nursery Lemoiue in 1889 (Nelson, pers. comm.). The plant was 
cultivated at Kew in 1896 (Royal Gardens 1896), in Ukraine in 1898 
(Kokhno 1986), and in the Botanical Garden in Darmstadt, Germany, in 
1900 (Purpus 1900). The Purpus article contains the earliest photograph 
of the species known to us. 

The seeds or plants fcr these early European introductions almost 

and garden there had long been receiving plant marerials collected by Rus- 
sian travellers in central and eastern Asia. Duplicates of these collections 
were sent to other major European botanical gardens (Bretschneider 1898). 
As many botanical gardens do, the one at St. Petersburg published annu- 
ally a list of seeds available; L. maackii first appeared m the garden's list, its 
Delectus seminum, in 1887 (Hortus Botanicus Imperialis Petropolitanus 

Within a few decades, Amur honeysuckle was growing in botanical gar- 
dens through much of Europe. Seeds of the plant eventually were offered in 
the seed lists of various European gardens for the first time in the following 
years: Cambridge, 1913 (Cambridge University Botanic Garden 1913); 
Oslo, 1917 (Universitet Botanske Have 1917); Dublin, 1919 (Royal Botanic 
Gardens, Dublin, 1919); Copenhagen, 1924 (Horto Universitatis 
Hauniensis 1924); Edinburgh, 1924 (Royal Botanic Garden, Edmburgh, 
1924); Amsterdam, 1929 Qardin Botanique de I'Umversite d'Amsterdam 
1929); and Pans, 1931 (Museum d'Histoire Naturelle 1931). 

From the Pacific to Western Europe: The English Role 

The earliest recorded observation of L. maackii by a European is not that 
of Maack but appears to be that of Robert Fortune, who in the mid 1840s 
collected the species in China. Where in China the specimen came from, 
either Amoy or somewhere in "northern China," has been a matter of de- 
bate. The specimen (two sheets at Kew) has but scant data: "A" and "34" 
(Fig. 2). Bretschneider (1894, 1898) concluded that the "A" stands for 
Amoy, which Fortune did indeed visit. The specimen may well have been 
collected in a garden because Fortune spent much time searching gardens 
for new plants to introduce to Europe. 

Several authors (e.g.. Anonymous 1929, 1934; Bean 1973; Thatcher 
1922; Wilson 1929) maintained that the first introduction of L. maackii 
into Great Britain was in 1900 by E.H. Wilson; it was one of the species he 
collected in China during his first trip there for the James Veitch Nursery. 
We were, however, unable to reconcile this date and method of introduc- 
tion with the statement in Bretschneider (1898) that the St. Petersburg 
Garden sent to the "greater botanical institutions in Europe and America, 
especially to Kew" (italics ours), seeds and plants and also duplicate speci- 
mens from the collections it received from central and eastern Asia. Be- 
cause L. maackii was growing at St. Petersburg since about 1880 and was 
first listed in the Garden's Delectus semtnum in 1887, we wondered why 
propagules of the plant had not been sent to Kew before 1900. After read- 
ing in Truelove (1917) that L. maackii was listed in "1894" in the Kew 
"Hand-List of Trees and Shrubs" we finally obtained a copy of that work 

LUKEN AND Thieret, Lonicera maackii 485 

(date actually 1896 for volume 2, the one of concern) and found that L. 
maackii is indeed listed there (Royal Gardens 1896). We suggest that seeds 
from St. Petersburg were sent to Kew some time before 1896 and that the 
plant or plants from those seeds languished, unheralded, among their con- 
geners in the garden. However, what might be called the "effective" intro- 
duction of L. maackit into Britain was apparently that in 1900 by Wilson 
for the Veitch nursery, which then extolled and disseminated it in Britain 
and elsewhere (Allan 1974). Once the Veitch propaganda machine was ac- 
tivated in its behalf, the plant received many notices in horticultural lit- 
erature, largely favorable until recently in North America. The company 
exhibited specimens of Amur honeysuckle at a meeting of the Royal Hor- 
ticultural Society in 1907 where they received an Award of Merit (Floral 
Committee 1908; see Anonymous {19151 for a drawing of L. maack'u made 
from the 1907 Veitch specimens.). A similar award was bestowed on the 
plant in 1915 (Floral Committee 1916). Amur honeysuckle was one of the 
few plants to which, until that time, such a double award had been made 
(Truelove 1917). Early mentions of the plant in continental European peri- 
odicals include one in the Belgian Tribune Hortkole, with a photograph sup- 
plied by Veitch (Anonymous 1909), and one in the French Revue Horticole, 
reporting introduction of the species into France apparently through the 
agency of Veitch (Mottet 1907). Veitch sent seeds of the plant to the United 
States Department of Agriculture (U.S.D.A.) as early as 1908 (U.S.D.A. 
From the Pacific and Western Europe to North America 

The earliest North American record of Amur honeysuckle we have lo- 
cated is in archives of the Dominion Arboretum, Ottawa: plants were re- 
ceived there in 1896 from Spaeth Nurseries in Germany (Cole, pers. comm.). 
The first U.S. record is in archives of the New York Botanical Garden: 
seeds of Amur honeysuckle from Russia were accessioned there in 1898 
(Riggs, pers. comm.) (Table 1). 

This first U.S. record of L. maackii came about through the agency of the 
then newly organized Section of Foreign Seed and Plant Introduction (S.PI.) 
of the U.S.D.A., which was mandated to procure, propagate, and distrib- 
ute new and valuable seeds and plants. In 1897 the U.S.D.A. dispatched 
Niels E. Hansen as an agricultural explorer to Russia in search of cold- 
hardy forage plants. The trip, Hansen's first for the U.S.D.A., extended 
from June 1897 to March 1898 (Hansen 1909; Taylor 1941). Unilaterally 
expanding his charge, Hansen sent about 9.30 accessions of forage, shrub, 
and tree seeds to Washington, DC, between December 1897 and June 1898 
(U.S.D.A. 1899a, 1899b). Some of the seeds were delivered before facili- 
ties were ready for their storage and dissemination (Fairchild 1938). 

486 SiDA 16(3) 1995 

Distribution of seeds received by the Section was started soon after they 
were received at Washington. One of the first recipients of seeds was the 
New York Botanical Garden: the "PIE" in the 1898 entry for L. maackii in 
the Garden archives indicated one of the first "Plant Introduction Experi- 
ments"— i.e., seed distributions— initiated by the S.RI. 

The seeds and plants imported by the S.P.I, were numbered consecu- 
tively starting in "Inventory No. 1," 1898. Hansen's collections are listed 
in the first two inventories (U.S.D.A. 1899a, 1899b). The data for number 
246 in Inventory I are "Lonkera maackii. From Russia. Received through 
Prof. N.E. Hansen, December, 1897." A similar entry, dated January 1898, 
is number 391 in this first inventory. Seeds of Amur honeysuckle were thus 
among the first few hundred accessions received by the S.P.I. 

The geographical origin of the seeds of L. maackii sent by Hansen is an 
intriguing mystery. According to the inventory data (U.S.D.A. 1899a, 
1899b), the sources of Hansen's collections seemed to have spanned much 
of Russia from St. Petersburg and Odessa to the Pacific. Origins of most of 
the seeds, including those of Amur honeysuckle, are given in broad terms, 
often simply "from Russia." More exact data are given for a few species: 
some came from "Sea Province {now Primorski Krai or Maritime Terri- 
tory], South Ussurie, Siberia" and some from "Amur." However, in spite of 
data indicating far eastern Russia, Hansen's 1897-1898 journey did not 
extend into that part of Asia. 

The Russian segment of his journey began and ended at St. Petersburg 
via Tashkent, Semipalatinsk, and Omsk (Taylor 1941). Apparently the far- 
thest east he travelled was when he visited Kuldja (or Kulja; also known as 
Gulja, Ining, and Yining), a Chinese city in western Sinkiang within ca. 50 
miles from the Russian border. (A rather difficult-to-interpret map show- 
ing the routes of Hansen's several Asiatic trips was published in Hansen 

Even though the exact western Chinese range of L. maackii is uncertain, 
the species is not known to occur in that small portion of China visited by 
Hansen in 1897-1898 (Hsu and Wang 1988). The seeds he sent to Wash- 
ington, then, must have come from some botanical garden, forestry sta- 

such establishments whenever he had the opportunity. For example, in 
August 1897 he was at the St. Petersburg Garden (U.S.D.A. 1899b); seeds 
of Amur honeysuckle certainly were available to him there from the stock 
maintained by the Garden for exchange. 

Thinking that Hansen's seeds marked simply "from Russia" might have 
been obtained from St. Petersburg, we obtained a photocopy of the garden's 
1899 Delectus seminum (the Delectm for 1897— the year of Hansen's visit 
there— and for 1898 were not available to us). The list (Hortus Botanicus 

LuKEN AND Thieret, Loniccra maackii 487 

Imperialis Petropolitanus 1899) contains a most impressive number of en- 
tries — some 3000 of them; some of the species represented among Hansen's 
seeds are in the Delectus, but most are not. Lonkera maackii is there, as it is 
in the 1887 Delectus. St. Petersburg, then, could have been the source of 
Hansen's Amur honeysuckle seeds, but they could have come from some 

After Hansen introduced Amur honeysuckle, the U.S.D.A. imported it 
from foreign countries and released it in the U.S. at least eight times through 
1927 (Table 1). Some of the introductions were from British botanical gar- 
dens; others were collected from native habitats in Manchuria by U.S.D.A. 
employees. The success of this introduction effort was indicated by the fact 
that in 1931 Amur honeysuckle was available from at least eight commer- 
cial nurseries throughout the U.S. (Partington 1931). The history of intro- 
duction published by the U.S.D.A. indicates that plants of the honeysuckle 
now naturalized throughout eastern U.S. represent a mixture of genotypes 
of diverse origins. 

Beginning in the 1960s and culminating in five official introductions 
up to 1984, the U.S.D.A. Soil Conservation Service (S.C.S) sponsored a 
program to develop improved cultivars of Amur honeysuckle. It was hoped 
that these cultivars would further traditional goals of the S.C.S. : soil stabi- 
lization/reclamation and wildlife-habitat improvement. From plants already 
naturalized in various parts of the U.S., genotypes were selected for more 
abundant fruit production, propagated vegetatively, and then cultivated in 
seed production blocks at various plant materials centers around the coun- 
try (Sharp and Belcher 1981). Seeds were made available by request. The 
most successful of these cultivars is 'Rem-Red' (Lorenz et al. 1989). 

AMUR honeysuckle: escape and naturalization 
In the New World 

The earliest hint we have located of the plant's escape in the New World 
is in archives of the Morton Arboretum near Chicago, which mention its 
weedy tendencies: "weed in arboretum since 1924, when first brought in" 
(Swink, pers. comm.). This early hint of the plant's decline in favor at Morton 
has accelerated toward a fall, the current situation there being well stated 
by Swink and Wilhelm (1994): "It would be difficult to exaggerate the 
weedy potential of this shrub." Floyd Swink has remarked to us that the 
spread of L. maackii in the arboretum is "unbelievable ... it would take a 
full-time worker to keep Amur honeysuckle and Rhamnus cathartica under 

A host of full-time workers would be required to keep Amur honey- 
suckle "under control" in the Greater Cincinnati region (including far north- 
ern Kentucky) from which the plant was first reported for Ohio by E. Lucy 

Braun (1961) only from Hamilton County, where it was "becoming abun- 
dant in pastures and woodlands." (As of October 1994 specimens have been 
collected in 34 Ohio counties {Trisel, pers. comm.}). In Greater Cincinnati 
the plant is now omnipresent, being by far the area's commonest shrub, 
native or alien. Efforts — in part thwarted by birds — are being made by 
various governmental agencies to eliminate the species from woodlands 
and other sites. The plant's establishment has been little short of phenom- 
enal. The species is ubiquitous, and often abundant, on open slopes and in 
fencerows, pastures, prairies, thin woods, woodland borders, road rights- 
of-way, railroad yards, and waste places. When the junior author moved 
into his home in Alexandria (Campbell County, KY) in 1973, no Amur 
honeysuckle was on the property. Now hundreds of individuals are there. 
We conducted a survey of selected botanical gardens and arboreta in the 
eastern United States and in eastern and western Canada. Although many 
botanists and arborists who responded noted that L. maackii was natural- 
ized, the species was considered a problem weed only in the following lo- 
calities: National Arboretum, DC; Morton Arboretum, IL; Butler Univer- 
sity, IN; Bernheim Forest, KY; Matthei Botanical Gardens, MI; W.J. Beal 
Botanic Gardens, MI; Shaw Arboretum, MO; Morris Arboretum, PA; Core 
Arboretum, WV. The species may not be winter hardy at Edmonton and 
Montreal. Non-cultivated plants of Amur honeysuckle are currently known 
in at least 24 states of the eastern U.S. (Trisel and Gorchov 1994) and in 
Ontario (Pringle 1973). 

The spread of Amur honeysuckle beyond the points of introduction is 
clearly facilitated by bird dispersal (Ingold and Craycraft 1983). Dirr (1990) 
noted that "birds deposit the seeds in old shrub borders, hedges, wasteland 
and before one knows it, Amur honeysuckle has taken over." Seed produc- 
tion is so prodigious, however, that most seeds simply fall to the soil. Such 
gravity dispersal, plus rainwash at least on slopes, is sufficient to allow 
seedling establishment at edges of existing colonies or mdividuals (Luken 
and Goessling 1995). Soil seedbanks as high as 1100 seeds/m^ have been 
recorded (Luken and Mattimiro 1991). 

Despite the published record of the plant's invasiveness, a U.S.D.A.- 
S.C.S. leaflet (Anonymous 1977) described Amur honeysuckle as a useful 
specimen plant for borders, hedges, and screens. Then followed the under- 
statement that the species "may spread into old fields, fence rows and open 
woodland." Other sources (Anonymous 1980; Sharp 1970) asserted that 
the shrub has "no objectionable features." Amur honeysuckle is, unwisely, 
still recommended (Lorenz et al. 1989) and commercially available (e.g., 
Southmeadow Fruit Gardens 1994). 

Not until the 1960s, after the shrub achieved some notoriety as a "weed" 
of concern in the U.S., did ecological studies of Amur honeysuckle begin. 
The data base on the species has been greatly augmented through research 
on population structure, productivity, interaction with herbivores, and com- 
munity dynamics (Ingold and Craycraft 1983; Luken 1988; Luken and 
Goessling 1995; Williams et al. 1992). As yet, no definitive study has been 
undertaken to determine if invasion of Amur honeysuckle is directly linked 
to local changes in native floras. 

Because of conservation goals calling for native species only, numerous 
prescriptions are now available for "control" of this species and for elimi- 
nating It from natural areas (Nyboer 1992). What is probably the nadir in 
the fall of Amur honeysuckle was reached in 1989 when the Illinois De- 
partment of Conservation adopted a policy mandating that no uses of the 
plant are acceptable in that state (Harty 1993). 

Even though L. maackii has been cultivated in Europe longer than in 

weediness there. One possible explanation is that fruit production by the 
shrub in at least western Europe, especially in England, seems to be less 
predictable than it is in eastern North America. The original report of 
flowering of Amur honeysuckle in eastern Europe (Kegel 1884) mentioned 
the fruit, but early western European accounts described flowers only — e.g., 
Belgium (Anonymous 1909), France (Mottet 1907), Germany (Purpus 1900; 
Schneider 1911), and Great Britain (Anonymous 1907a, 1907b, 1915). 

490 SiDA 16(3) 1995 

An early report from Germany (Dippel 1 889) noted the lack of fruit devel- 
opment. Not until 20 years after the species' introduction into Great Brit- 
ain were the fruits described in British horticultural literature (e.g., Anony- 
mous 1917). The fruitmg habits of L. maackii m England were singled out 
for comment in two articles. The first cautioned that "no shrubby Honey- 
suckles can be depended on to fruit with the same constant profusion in 
our English climate as they do in the United States, where . . . gardeners do 
not suffer from frosts after winter is over" (Thatcher 1922). The second 
mentioned that "as a fruiting shrub ... its merits are not so well known, 
but, apparently, in warm seasons and on certain soils it fruits abundantly 
and becomes a highly attractive plant in October" (Anonymous 1934). 
Another explanation may be the fact that L. maackii apparently is grown in 
Europe largely on estates or in botanical gardens where weediness is often 
simply not tolerated, any volunteer plants being quickly removed. Fur- 
ther, climatic differences between western Europe and eastern Asia/eastern 
North America may be such that potential for naturalization is suppressed. 

In Eastern Asia 

In its Asiatic range, L. maackii occurs in "mixed forests" (North Korea; 
Kolbek and Kucera 1989); edges of montane deciduous forests, sometimes 
on calcareous rocks (Japan; Hara 1983; Iwatsuki et al. 1993); in oak- and 
elm-woodland (Amurland; Herder 1878; Ruprecht 1857); in xerophytic 
forests dominated by spinous-leaved oaks (northwestern Yunnan; Forrest 
1915); in forests oi Ab'm delavayi, Picea yunnanensis and other Picea, Tsuga 
yunnanemts, and species o^ Acer, Prunus, Pyrus, Sorhus, and Tilia (northwest- 
ern Yunnan; Forrest 1916); and forests oi Liriodendron cbinense (western China; 
Wilson 1913). In 1994 at Changbai Mountain Forest Research Station in 
northeastern China, the first author (JOL) found Amur honeysuckle grow- 
ing exclusively in frequently disturbed floodplain forests with Acer mono, 
Fraxinus mandschurka,Juglans mandshurka, Phellodendron amurense, Populm 
davtdiana, Prunuspadm, Syringa amurensis, and Ulmus propinqua. In the lower 
elevation montane forest at Changbai, the species grew with Lartxprincips, 
Quercus liaotungensis, and Tilia mandshurica. According to Wang (1961) L. 
maackii was part of the undergrowth of shrubs in the "Montane-boreal Co- 
niferous Forest of the Northeastern Province," which is characterized by 
Abies holophylla, A. nephrolepis, Picea jezoens is, P. obovata, and species ofLarix, 
Pinus, and Taxus. Forrest ( 1 9 1 5 , 1 9 1 6) and Wilson (1913) listed other woody 
associates in China: Amelanchier asiatica, Berberis sp., Corylus sp., Crataegus 
cuneata, Deutzia sp., Diervilia [Weigela)japonica, Indigoferapendula, Ligustrum 
ionandrum, L. Henry t, L. ligustrina ior:m2i yunnanensis, Lonicera xerocalyx, Phila- 
delphus sp.,Rhamnus sp., Rosa sericea, Styrax hemsleyanus, Symplocos crataegoides , 

..„,„, and Wikstroemia sp. Woeikoff (1941) remar 
iber" of species ofLonicera, including L. ?naack.ii, 

In North America 

As is the case with many successful introduced species, L. maackii thrives 
in communities with histories of human disturbance. Specifically, urban or 
exurban forests with histories of fragmentation, cattle grazing, or wood- 
cutting are often heavily invaded. In such sites in northern Kentucky/south- 
western Ohio L. maackii forms a dense understory and associates with Acer 
saccharum, Celtis occidentalism Fraxinus americana, Gleditsia triacanthos, Madura 
pomifera, Prunus serotina, Quercus rubra, Robinia pseudoacacia, and Ulmus rubra 
(Luken 1988, 1990; Williams et al. 1992; Yost et al. 1991). In large, closed 
canopy forests, L. maackii may be relegated to forest edges, suggesting 
moderate shade intolerance. 

The plants are fully capable of thriving in full-sun environments. Pro- 
ductivity in open sites is higher than in forests (Luken 1988); open-grown 
plants also show strong resilience when cut repeatedly (Luken and Mattimiro 
1991). Open sites heavily invaded by L. maackii may, after invasion, be 
converted to a scrub-type community. The understory in this scrub is often 
dominated by the grasses Festuca arundinacea and Poapratensis and by intro- 
duced herbaceous species such as Coromlla vana and Melilotus alba (Luken 
and Thieret 1987; McClain and Anderson 1990). 


Lonicera maackii is part of many publications on the flora of Japan, China, 
Korea, and far eastern Russia. The information available in these varies 
from mere mention in checklists to accounts with descriptions, ecological 
and morphological notes, and geographical data. Some works of the latter 
kind are, for Japan, those by Kara (1983), Iwatsuki et al. (1993), Kurata 
(1971), Nakai (1921b), and Ohwi (1965); for China, those by Hao (1934), 
Hsu and Wang (1984, 1988), Komarov (1907), and Noda (1971); for Ko- 
rea, those by Kolbek and Kucera (1989), Lee (1989), and Nakai (1909, 
1911, 1921a); and for Russia, those by Charkevicz (1987), Maximowicz 
(1878), Penkovsky (1901), Poyarkova (1958), and VolT (1899). The earli- 
est reports known to us from countries in the species' range are the follow- 
ing: Russia (Ruprecht 1857), China (Maximowicz 1878), Japan 
(Maximowicz 1878), and Korea (Palibin 1898). 

North America 

prisingly slow. Examples follow. 

Although collected in Maryland as early as 1937, Amur honeysuckle is 
absent from Gray's manual of botany (Fernald 1950) and from The new Bntton 
and Brown illustrated flora of the northeastern United States and adjacent Canada 
(Gleason 1952). Two decades later, m the Gleason and Cronquist (1963) 
Manual, it received passing mention — "it is becoming established in 

Maryland" — but was not included in the key to species of Lomcera. By the 
1991 edition of the Manual (Gleason and Ctonquist 1991) it had been 
admitted and placed in the key as a full-fledged member of the flora: "es- 
caped and becoming naturalized in our range, as in N.Y., Md., Ky., and 
Oh." (The species was not listed for New York just 5 years before [Mitchell 
1986], even though herbarium records for that state indicate the presence 
of Amur honeysuckle there as early as 1954 [Trisel and Gorchov 1994}). 

Amur honeysuckle was collected as early as 1966 in Virginia (Trisel and 
Gorchov 1994), but it is missing from the 1981 atlas of the Virginia flora 
(HarviU et al. 1981). In the 1992 edition (Harvill et al. 1992) it is listed 
from seven counties. The plant is "taking over people's back yards" in parts 
of Virginia (Luken, personal observation). 

Some field guides, too, have been slow in recognizing the establishment of 
Amur honeysuckle in North America. For example, the plant is not to be 
seen in Petrides' field guide (Petndes 1986) to "all" trees, shrubs, and woody 
vines in the northeastern and central United States and adjacent Canada. 

.- honeysuckle, Maack's honeysuckle, 
or even (through misinterpretation of the specific epithet), "Maacky's" hon- 
eysuckle (Darnell 1930), Amur honeysuckle is a robust, upright, multi- 
stemmed, deciduous shrub that can grow to 6 m tall, with stems to 15 cm 
in diameter near the base, and can attain a spread of 9 m (Doney 1947). 
The bark of the largest stems, longitudinally fissured, is dark gray to gray- 
ish brown. Long shoots can grow as much as 1.2 m in their first year and 
can bear some flowers. Short shoots develop from axils of long shoots; they 
flower freely. The branchlets are hollow. Dark green and lightly pubescent, 
the leaf blades average about 7 cm long; on fast-growing shoots they may 
reach 15 cm. Their acuminate apex is one of the species' distinguishing 
features. Distinctive, too, is the early leafing out of the species before that 
of associated deciduous-leaved plants and the persistence of the leaves in 
fall, sometimes into December (a few even into January) in central U.S. 

The paired, axillary flowers (Fig. 3), produced in profusion especially on 
short shoots, are borne on peduncles 1.5 to 5 mm long (shorter than the 
subtending petioles); variation in peduncle length can be seen on a single 
branch. Five-lobed, the calyx is 2 to 3 mm long and eventually deciduous. 
Like those of some other honeysuckles, the white to pink corollas, 1.7 to 
2.5 cm long, are two-lipped, the upper lip four-lobed, the lower lip entire. 
In age, the corollas become a dull yellow, a color change seen also in certain 
other species of Lomcera, e.g., L. japonka, L. morrowii, and L. tatanca. 

494 SiDA 16(3)1995 

Characteristics of the flowers make Amur honeysuckle a good bee plant. 
Unlike those of some other species o{ Lomcera, the "corolla tubes are short 
enough for honey bees to work successfully, and good nectar secretion takes 
place in cool, damp weather as well as under hot, dry conditions" (Clark 

The twinned berries, admittedly one of the glories of the species, mature 
in fall. Ranging in diameter from 3.8 to 8.5 mm, they are glossy, translu- 
cent red (sometimes with a slight orange cast), and ± globose to somewhat 
ovoid or ellipsoid. To us they look much like red currants {Ribes). In mild 
seasons the fruits may persist, on leafless plants, until nearly Christmas in 
the Northern Kentucky/Cincinnati area. 

The fruits of some honeysuckles have been reported to be toxic. In a 
study by Leveau et al. (1977) the fruits of eight species, including L. maackii, 
were found to "contain a small quantity of alkaloids, but saponosides seem 
to be responsible of (i.e., for} toxicity"; "immature fruits are more toxic 
than mature fruits" and "pericarps are much more toxic than seeds." Frohne 
and Pfander (1984) suggested that, for problems to develop, about 30 fruits 
would have to be eaten, which is not likely to be done by people with 
functional taste buds — the fruits are exceedingly bitter. 

A chromosome count o{2n = 18 was published for Lonkera maackii var. 
maackii and for L. maackii var. erubescens (Janaki Ammal and Saunders 1953). 

Horticultural Value 

Its serious drawbacks in North America notwithstanding, Amur honey- 
suckle is a most handsome shrub, whether in flower or in fruit. It has been 
much extolled for horticultural purposes, as the following quotes attest. 
"Among all eastern Asiatic species {o^Lomcera} L. maackti is the most beau- 
tiful" (Regel 1884). "There are few of the bush honeysuckles more beauti- 
ful than this variety" (Anonymous 1917). "... among the three best bush 
honeysuckles for the pleasure grounds and shrubbery borders" (Anonymous 
1924). "... one of the most all-round beautiful members of an indispens- 
able family" (Wilson 1929). 

Its attractive foliage was singled out for special mention by some au- 
thors (e.g., Beezley 1939; Henry 1932; Purpus 1900). Alexander (1944) 
wrote that its "dark green foliage remains green until Christmas, thus ri- 
valling some of the hollies in outdoor effect." 

The fruiting habits of the plant have often been mentioned quite favor- 
ably: "Since It holds it fruit late into the winter it is a valuable addition to 
garden beauty when that beauty is limited" (Wilson 1925). "While nearly 
every other shrub is bare [L. maackii^ stands out with its mantle of green 
studded with red berries strong and defiant" (Wilson 1925, 1928). 

We assume that L. maackii has, by now, been widely distributed through- 

out the temperate horticultural world, even though we made no special 
effort to determine in what areas it is now grown. We did note, however, 
that It is cultivated in Italy (Cocker 1935), Japan (Kurata 1971), and New 
Zealand (Cook 1949). We have seen no published hints of the plant's weedi- 
ness in any area other than North America. 

1 Lonkera maackti has been controversial. Most 
authors cite Maximowicz as having made the combination Lonkera maackii 
in Prtmittae florae Aniurenm (Maximowicz 1859), but he did not validly 
publish such a new combination there. The binomial Lomcera maackii (Rupr.) 
Herder was validly published in 1864 (Herder 1864). 

The first infraspecific taxon within L. maackti was described by Rehder 
(1903): L. maackii {otmin podocarpa Franchet ex Rehder. A plant of western 
and central China (Sargent 1913; Schneider 1911) introduced by Wilson 
in 1900 (Farrington 1931), it was said to differ from the typical form as 
follows: "Has the ovaries, together with the bractlets, on a short, stalk-like 
elongation raised above the bracts which gives the impression of a some- 
what abnormal form, though it occurs in most Chinese specimens." Ten 
years later Rehder (1913) adopted varietal status for the taxon: L. maackii 
var. podocarpa (Franchet ex Rehder) Rehder, but in 1949 he returned to the 
status of forma (Rehder 1949a). Although a few authors have used the 
designation offorma (e.g., Anonymous 1929; Forrest 1916; Leveille 1915- 
1916) most consider the taxon to be a variety (e.g., Bailey 1916; Bean 
1914; Sargent 1913, 1922; Schneider 1911; Wilson 1913, 1917, 1925, 
1928;Wyman 1962). 

Variety podocarpa has been called "superior to the type" (Anonymous 
1924), "especially fine" (Hadden 1925), and "a better flowerer" (Dirr 1990). 
Bonstedt (1932), in contrast, said of it that "it is not so beautiful as the 
species." According to Wyman (1962), "The chances are that [L. maackii'\ 
and its variety podocarpa are badly mixed in nurseries." Hsu and Wang 
(1988) wrote that the feature by means of which Rehder originally distin- 
guished (ovma. podocarpa shows continuous variation in the species and thus 
no evidence has been found to support recognition of the form. We, too, 
have noted that the feature is quite variable, even on a single individual, 

A pink-flowered variant of L. maackii, described as forma erubescens by 
Rehder (1913), was said to have arisen from seeds collected by Wilson in 
China. The taxon is recognized as a form by some (e.g, Rehder 1949a; 
Sargent 1922) and as a variety by others (e.g., Alexander 1944; Bailey 1916; 
Hsu and Wang 1988; Royal Botanic Gardens, Kew 1934). We, however, 
do not recognize the variant at any rank because the corollas vary continu- 

4y6 SiDA 16(3) 1995 

ously from essentially pure white to a deep pink; all intergrades between 
these extremes can be found in a smgle population. Some of the pink indi- 
viduals we have seen are an excellent match for the colored illustration of 
"var. erubescens" in Addisonia (Alexander 1944); others, however, are much 
more deeply pigmented. 

According to Hara (1983), Japanese representatives of L. maackti "seem 
to differ slightly from the [East Siberian] plants ecophysiologically. When 
planted side by side in Tokyo, the East Siberian plants grow more vigor- 
ously bearing larger leaves with slightly impressed nerves, and flower and 
fruit much earlier, as compared with the Japanese plants." Earlier, Nakai 
(1938) had proposed a varietal name for Japanese plants but did not pub- 
lish it validly. And most recently Iwatsuki et al. (1993) did not recognize 
any infraspecific taxa for the Japanese representatives of the species. 

Accordingly, synonymy for Amur honeysuckle is as follows: 
Lonicera maackii (Ruprecht) Herder, Bull. Soc. Imp. Nat. Moscou 
37(1):204, 1864. 

Xylosteum nu.ackU Ruprecht, Bull. CI. Phys.-Math. Acad. Imp. Sci. St.-Petersbourg 

'; Capri foil ION ma 

'ackjj (Rupr.) Kuntze, Rev. Gen. Plant. 1 :274. 


het ex Rehder, Ann. Rept. Missouri Bot. Gard 


t (Franchet ex Rehder) Rehder, Mitt. Deutsch. 

)3. I9\ 

ackii forma eru/mcem Rehder, Mitt. Deutsch. 

>}. 1915; L. ma. 

tckn var. erukscens (Rehder) Rehder in Bailey, , 

:1910. 1916. 

::lusion: from 


The ascent of Amur honeysuckle began when the species was introduced 
from eastern Asia to St. Petersburg, where it first flowered in Europe. Be- 
cause of its attractive flowers and fruits, it was soon disseminated via di- 
verse channels to other parts of Europe and to North America where, as a 
new introduction, it won great praise. In the New World its decline in 
favor began once its weedy, aggressive, and invasive nature came to the 
fore. Though still touted — and even sold — by some for conservation or 
wildlife purposes, the shrub has fallen from grace rather recently when 
various authors warned that it should not be planted and a U.S. state even 
proscribed its use. 

The story of Amur honeysuckle recalls the stories of other Old World 
woody plants introduced into temperate eastern North America, only to 
become naturalized and often troublesome. Coming immediately to mind 
are Ailanthns altissima, Celastrus orbiculata, Elaeagnus umbellata, Lonicera 
japomca, Rhamnus cathartica, Rosa multiflora, and Ulmuspumila. One can but 
assume that the saga of introduction/naturalization, sometimes with un- 
fortunate consequences, will be a continuing one. 


We close our account of Amur honeysuckle with mention of the passing, 
ome 6 decades ago, of E.H. "Chinese" Wilson, the man responsible for the 
effective" introduction to Great Britain of the species — and of others that 
ave found their way mto western horticulture. On 15 October 1930 he 
nd his wife were killed in an automobile accident near Boston (Tozer 1994). 
it the double funeral service, the caskets were covered in floral tributes 
epresentmg some of Wilson's introductions (Briggs 1993). Among these 
ributes were fruiting branches o( Lomcera maackii. 

We thank the foUowmg individuals for aid during various stages of our 
project: Michael Agnes, David Boufford, David M. Brandenburg, Jean 
Cargill, Trevor Cole, Barney Lipscomb, E.C. Nelson, Becky Norris, Rich- 
ard Rabeler, Bruce K. Riggs, Joe Ruh, Richard Spjut, John Strother, Ronald 
Stuckey, Floyd Swink, Donald Trisel, and J. Wade Turner. Much use was 
made of collections in the Lloyd Library and the Missouri Botanical Gar- 
den. The Royal Botanic Gardens, Kew, sent the photograph used as Fig. 2. 
Financial support was provided by National Geographic Society, National 
Science Foundation, and Northern Kentucky University. 

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Amur honeysuckle {Lonicera maackii) during repeated clipping. Ecol. Applic. lilO'^ 

tucky roadside embankments. Trans. Kentucky Acad. Sci. 48:51-54. 
Maack, R. 1859. Puteshestvie na Amur [Journey on the Amur]. Tip. K. Vulfa 

Sr.Petersburg. (In Russian.) 
Mack, R.N. 1985. Invading 

Pages 127-142 /« J. White 

Academic Press, London. 
Maximowicz, C.J. 1859. Primitiae florae Amurensis. Mem. Acad. Imp. Sci. St. Petersbourg 

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Wissenschaften, Saint Petersburg.) 
1878. Diagnoses plantarum novarum Asiaticarum. II. Bull. Acad. Imp. 

Sci. Saint-Petersbourg, 3rd ser. 24:26-89. (Published also in Melanges Biol. Bull. Phys.- 

, T 1909. Flora Koreana. Pars prima. J. Coll. Sci. Imp. Univ. Tokyo ; 

1911. Flora Koreana. Pars secunda. J. Coll. Sci. Imp. Univ. : 

1921a. Flora sylvatica Koreana. Pars XI. Caprifoliaceae. The 

hosen, Seoul. (In Japanese and Latin.) 

Nicholson, G. 1888. llkistrated dictionary of gardening. Vol. 4. L. Upcott Gill, Lo 

NODA, M. 197 I . Flora of the n.-e. province (Manchuria) of China, No publisher indii 
no place indicated. (In Japanese and English.) 

Nyboer, R. 1992. Vegetation management guideline: bush honeysuckles— Tat; 
Morrow's, Belle, and Amur honeysuckle {Lonicera tatarica L., L. morrowii Gray, L. : 
Zabel, and L. maackii [Rupr.] Maxim.). Nat. Areas J. 12:218-219. 

Ohwi, J. 1965. Flora of Japan. Smithsonian Institution, Washington, DC. 

ActaHorti Petrop. 17(1):1-128. 

,UKEN AND Thieret, Lonicera maackii 501 

'ENKOVSKY, V.M. 1901. Derev'ia i kustarmki [Trees and shrubs]. Vol. 3. O.D. Khodshinoy, 
Kherson. (In Russian.) 

'ETRiDES, G.A. 1986. A field guide to trees and shrubs. 2nd ed. Houghton Mifflin, Boston. 

'OYARKOVA, A.I. 1958. Caprifoliaceae Vent. Pages 419-584 tn Flora URSS 23:419-584. 
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>iiRPUs, A. 1900. Lonicera maackt [sic] Maxim. Gartenwelt 4:235. 

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1913. Neue Oder kritische Geholze. Mitt. Deutsch. Dendrol. Ges. 191 3:254- 

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Vo!'i ^TTf 


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1991. The vegetation c 


Committee on Managing Global Genetic Resources. 1993. Managing 
Global Genetic Resources. Agricultural Crop Issues and Poli- 
cies. (ISBN 0-309-04430-8, hbk.). National Academy of Sciences 
Press, 2101 Constitution Avenue, N.W., Washington, D.C. 20418, 
(202) 334-3180. $49.95. 449 pp., 6x9. 

"Managing Global Genetic Resources. Agricultural Crop Issues and Policies examines the struc- 

legal, economic, and political issues that surround management and use. The Exe 
Summary, with the committee's major recommendations; an Overview, which intn 
the subject for those readers without a background in genetic conservation; and twc 
Part one addresses basic science issues and entails Chapters 1 to 10. Part two ad< 
policy issues and entails Chapters 1 1 to 15." The 15 chapters are: 1) Genetic Vulner 
and Crop Diversity; 2) Crop Diversity: Institutional Responses; 3) In Situ Conserva 
Genetic Resources; 4) The Science of Collecting Genetic Resources; 5) The Scie 
JManaging Genetic Resources; 6) Using Genetic Resources; 7) Biotechnoiog 
Germplasm Conservation; 8) Documentation of Genetic Resources; 9) The Conser 
of Genetic Stock Collections; 1 0) The Genetic Resources of Microorganisms; 11 ) Exc 
of Genetic Resources: Quarantine; 1 2) Exchange of Genetic Resources: Proprietary B 
13) Genetic Resources: Assessing Economic Value, 140 Conflicts Over Ownership, 
agement, and Use; and 15) National and International Programs. 

Primack, Richard B., 1993. Essentials of Conservation Biology. (1 

0-87893-722-6, hbk.). Sinauer Associates, Inc., Sunderland, M 
chusetts 01375, U.S.A. 564 pp. 

Diology. The text ope 

ns with 

1 concise c 


;ions of 

the origins of the disc 


ding chap 

ters re 

:view th. 

e applications of conse 

^ principles to problei 




: of habitat and remnai 

11 as human dominate 

be restored. 

^Conservation Biology 

will be 

useful in 



te introductory course 


g worL 

dwide cas< 


iples in 

the narratives as well ; 

The frequent figures 

are clear and we 


illustrate the narrativ 

logy, who usually represent a 





Botanical Research Institute of Texas 

^09 Pecan Street 

Fort Worth, TX 76102-4060, U.S.A. 

P. 0. Box 61 

San Gregorio, CA 94074, U.S.A. 

The genus Clusia sections Criuvopsis and Brachystemon are revised, in preparatio 
reatment of the Clusiaceae for Flora of the Venezuelan Guayana and Flora de Colomb'u 
ilified descriptions, keys to the species and lectotypifications are provided for each s 

•mazonka and C. martiana. Clusia oedematopoidea is reduced to synonymy under C. amc 
nd C. uleana under C.marttana. In Clusia section Brachystemon, 3 taxa are recogni 
vhich one, Clusta aymardii, is described as new, illustrated, and its phylogenetic re 
hips are discussed. Annplified descriptions and discussions of phylogenetic relatic 
.re provided for Clusia spathulaefolia and C. penduliflora. 


The genus Clusia Lii 

chystemon Se presenta 

n descripciones act 

ualizadas para las dos 

■susespecies,y select 


anoce tres especies en 

are, se describe coi 

no especie nueva, se 

entesco. Se provee d 

adas, ilustraciones y 

filogeneticas para Clusta amazomca,y C. i 


L C. uleana bajo C. 

martiana. In seccion 


ales, C.^p«Wn sec 

; su parentesco. Tamb 

•en, se provee descr 

ipciones ampliadas, y 

i filogeneticas para C. 

spathulaefolia y C. 



inaeus contains o^ 

.er 250 species ( 

with 336 names), 

: the Neotropics, ^ 

with two, C. rose 

'rfjacquin, and C. 

■ cultivated throu 

ighout the tropi 

ics. Despite taxo- 

506 Sjda 16(3) 1995 

nomic overdescription, it is estimated that there are at least an additional 
40 as yet undescribed species in the Andes of Colombia, Ecuador, Peru and 
Bolivia, particularly in the premontane, montane (including ceja de selva), 
ubparamo, paramo and jalca habitats. The genus is easily recognized by 
fruits, with few to numerous stamens, 
e latex. Characters traditionally used to 
dehne the sections of the genus include the structure of the androecium in 
staminate flowers, and corresponding structure of the stigmas and 
staminodes of the pistillate. The last comprehensive revision was that of 
Vesque (1893), in which Engler's (1888) classification was simplified. The 

nate, pistillate, bisexual (Hammel 1986) and apomictic (Hammel 1986; 
Maguire 1976) individuals known. In addition, I have seen staminate flowers 
on reiterative (sensu Halle et al. 1978) shoots of otherwise functionally 
pistillate plants in Climapimlla Steyermark and in C. grandiflora Splitgerber, 
in the Pakaraima Mountains of Guyana and on the Chimanta Massif com- 
plex of tepuis in the eastern portion of the state of Bolivar, Venezuela. 

While preparing a diagnostic treatment of the genus for the Flora of the 
Venezuelan Guayana and a more comprehensive one for Vlora de Colombia, 
sections Criuvopm and Brachystemon were revised. These two sections form 
a natural group, closely related to section Criuva Bentham. Engler (1893) 
considered both sections Criuvopm and Brachystemon to be subsections within 
subgenus Criuva sensu Bentham (1862). In fact, this study grew out of the 
difficulty in distinguishing young plants oiClusia {Brachystemon) pendu 1 1 flora 
from C. {Crwvopsis) marttana. Because the formats of the floristic treat- 
ments in preparation do not allow for complete descriptions and synonymy, 
the present treatment is intended to provide data for the groups over their 
entire range, until a more comprehensive revision is prepared. 


isia L. section Criuvopsis Planchon and Trian 
13:322. 1860. Lectotype Specii^s (here designated 
and Triana, Ann. Sa. Nat., Ser. 4 13:358. I860. 

a, Ann. Sci. Nat., Ser. 4 

lusia section Cnuva Bentham and Hooker, subsection Cri 
Engler, Fl. Bras. 12(1):401. 1888. J. Vesque in A. DC a 

uvopus (Planchon and Triana) 
nd DC, Monogr. Phan. 8:77. 

lusia subgenus Criuva (Bentham and Hooker) Engler, sec, 

cion Criuvopus (Planchon and 

Inana) hngler, Nat. Pflanzenfam. 3(6):225. 1895. 
Glabrous lianas or hemiepiphytes; latex white to translucent. Leaves petio- 
late. Inflorescence a terminal panicle, the flowers cymose; secondary inflo- 
rescence bracts 2, coriaceous; bracteoles 4, decussate, coriaceous. Perianth 
spreading; sepals 4-5, char taceous, opposite the petals; petals 5, contorted, 

PiPOi.Y AND Grai-t, Synopsis of Clusia 50 / 

very thickly carnose; androphore pentagonal, stamens numerous, the fila- 
ments short, flat, apically free, connate at the base, fleshy, often thickened 
at the base; anthers basifixed, muticous, equalling or slightly longer than 
the filaments, truncate to emarginate apically, dehiscent by longitudinal 
slits; pistiUode absent, the central receptacle resiniferous or not. Pistillate 
inflorescence and flowers as in staminate, but staminodia 5, free, linear, 
with vestigial anthers; carpels 5, stigmas thin, elliptic to oblong, peltate; 
ovules horizontally ascending;. Fruit a capsule, ovoid to oblongoid, stigma 
and styles persistent. 

Distribution. — Three species, distributed from Panama to Colombia, 
Venezuela, Ecuador, Peru, Bolivia and Amazonian Brazil. 

Clusia section Criuvopsis appears to be most closely related to C. section 
Brachystemon by virtue of its short, thick filaments of the fertile stamens, 
sessile stigmas, and linear anthers deshiscent by longitudinal slits. How- 
ever, section Criuvopsis may be easily distinguished from section Brachystemon 
by the flat filaments connate basally, and very thin inflorescence rachises. 





Leaf blades ( 

)blong, bulh 

ite; sepals 

4; petals 5 

,; anthers rou 

nded to slight 

emarginate i 

basally .... 


.Leaf blades c 

)bovate to ell 

iptic, smooth; sepals 

,5, petals 5; a 


2. Leaf base 

us. adaxial la 

adaxial ve 

d submargin. 

l1 collecting v 



uex canals hig 

)us, adaxial v 

enation in. 


IS, submargin 



iscern.ble; s 



: shrub, at t 

imes appear; 

. Clusia ar 


. Pipoly, 

sp. nov. 

(Fig. 1) 

Scandent lianous shrub, to 23 m long, 4 cm diam. glabrous; branchlets 
tetragonal in cross section, squarrose, 3-5 mm diam., bark of apical por- 
tions rufous, appearing scaly, checking transversely, gray at maturity, rug- 
ose, latex white, copious. Leaves decussate; blades coriaceous, oblong, (8)13- 
18 cm long, (2.5-)3-3.5(-4) cm wide, apex abruptly acuminate to rostrate, 
base widely rounded to truncate, deeply bullate, nitid above at first, then 
pallid above and below at maturity, midrib deeply impressed above, promi- 
nently raised below, secondary veins 17-32 pairs, perpendicular to midrib, 
deeply impressed above, prominently raised below, united by a prominent 

showing the oblong, bullate leav 
;. C. Quadrate pedicel. D. Outer s 
pentagonal morphology and apically r 

PiPOLY AND Graff, Synopsis of Clusia 509 

submarginal collecting vein, latex canals linear, conspicuous, margin revo- 
lute, entire; petioles deeply canaliculate, 2.5-2.8 mm long, glabrous. Stami- 
nate inflorescence terminal, erect, pyramidally paniculate, 7-9 cm long 
and 5-7 cm wide in bud, the branches cymose, flowers in 3's; peduncle 
3-5-5 cm long, subtended by two reduced leaves similar to vegetative leaves 
but 3-5 cm long, 1.4-1.8 cm wide; secondary inflorescence bracts carnose, 
depressed-ovate, 1.3-1.6 mm long, 1.6-1.8 mm wide, apex rounded, me- 
dially carinate, the margin opaque, not scarious, entire; pedicel squarrose, 
2.8-4.2 mm long; bracteoles four, 2 decussate pairs, carnose, depressed- 
ovate to suborbicular, apex obtuse, carinate medially, the keel callose, hya- 
line, the margin entire, glabrous, the outer 2-2.5 mm long, 2.3-2.6 mm 
wide, the inner 2.5-3 mm long and wide. Staminate flower buds cream; 
outer sepals 2, opposite, coriaceous, orbicular, 2.3-6 mm long and wide, 
apex very widely rounded, cucullate, hyaline except apically, the margin 

pals but reddish, hyaline apically; petals 5, contorted, carnose, oblong, 0.8- 
1.1 mm long, 0.4-0.6 mm wide in bud; androphore convex, pentagonal as 

0.6-0.8 mm long, the anthers linear. 
rounded apically and basally; pistillod 

Typi.: COLOMBIA. Amazonas: Araracuara, along Rio Caqueta, right bank, 3 km above 
Sumaeta, ()°36'S, 72°]0'W, riparian floodplain, 200-300 m, 30 Sep 1990 (stam. fl bud), 
E. Alvarez. A.C. l.ondom. A. Rodriguez, & F. Moreno 116 (holotype: COAH!; isotype: NY!). 

Distribution. — Known only from the type. 

Ecology and conservation status. — Clusia araracuarae occurs in Igapo veg- 
etation (Prance 1979), forming mats in canopies of trees along black water 
river margins. The area in which this species occurs is subject to extreme 
disturbance owing to forest destruction, largely for firewood. Araracuara is 
a significant population center and busy port on the Rfo Caqueta, and there- 
fore, this species should be considered threatened. 

The long inflorescences, lianous habit, leaves with prominent secondary 
venation and submarginal collecting vein and obtuse to broadly rounded 
bases, indicate that Clusia araracuarae is most closely related to Clusia amazonica. 
However, the bullate leaf blades, fewer sepals, and anthers that are rounded 
to emarginate apically and basally clearly set C. araracuarae apart. 

2. Clusia amazonica Planchon and Triana, Ann. Sci. Nat., Ser. 4 13:358. 
I860. (Fig. 2). Type. BRAZIL. Amazonas: near Panure along the Rio Uaupes, 
Oct 1852-Jan 1853 (stam. fl),/?. Spruce 287 8 {ho\.otyv^: P!; NYNeg. 5033; isotype: 
P!;NYNeg. 5034). 

. 1978. Type. PANAMA. 

Duke 1355 

: MO!) 

Liana; branchlets tetragonal in cross section, squarrose, (2.5-)3-5.5(-7) 
mm diam., bark of apical portions rufous, appearing scaly, checking trans- 
versely and exfoliate, beige at maturity, rugose; latex white, copious. Leaves 
decussate; blades coriaceous, oblong, elliptic or rarely obovate, (8.0-)10.5- 
20.0(-26.0) cm long, (3.5-)4.0-4.6-10.5(-12.2) cm wide, apex broadly 
rounded, with an abruptly short-acuminate tip, the acumen 0.5-1 cm, base 
obtuse to broadly rounded, not decurrent on the petiole, smooth, sordid 
above, pallid below, latex canals varying in conspicuousness above, but never 
readily visible without maginification below, midrib prominently raised 
above and below, secondary veins numerous, prominently raised above, 
slightly raised below, united by a fine submarginal collecting vein, the mar- 

PiPOLY AND Grafp, Synopsis of Clusia 5 1 1 

gin subrevolute, entire, glabrous; petioles canaliculate, (2.1-)2.7^.0 (-5.8) 
cm long. Inflorescence terminal, erect, pyramidally paniculate, (6-) 12-24 
cm long, 6-10 cm wide at maturity, the flowers cymose, 3 per cyme; pe- 
duncle (2-)4-8(-11.5) cm long, subtended by normal leaves; secondary 
bracts coriaceous, very widely ovate, 1.5-2.5 mm long, 1.5-2 mm wide, 
apex obtuse to rounded, carinate, the margin hyaline, scarious, entire. Stami- 
nate inflorescence with tetragonal pedicels, 3-5 mm long; bracteoles 4, decus- 
sate, the outer 2 coriaceous, very widely ovate, 1.5-2.5 mm long, 1.5-2 
mm wide, apex obtuse to rounded, carinate, the margin hyaline, scarious, 
entire, the inner 2 suborbicular to orbicular, 2.5-3 cm long and wide, apex 
rounded, flat, the margins entire, hyaline, scarious. Staminate flowers white; 
sepals 5, contorted, membranaceous, brittle, suborbicular to orbicular, 5-6 
mm long and wide, apex broadly rounded, translucent, densely lineate, the 
margin hyaline, scarious; petals 5, contorted, carnose, somewhat cucuUate, 
oblong, 3-5-4 mm long, 2.5-3 mm wide, apex obtuse to rounded, some- 
what cucuUate, densely lineate, the margin hyaline; androphore flat or 
slightly concave, pentagonal, the stamens numerous, free, 2.5-3.5(-3.7) 
mm long at maturity, the filaments short, flattened, ca. 1-1.2 mm long, 
the anthers linear, 1.5-2.5 mm long, 0.7-1 mm wide, apex truncate, base 
truncate, dehiscent by wide longitudinal slits, abruptly expanding at base 
to filament, the connective flat, darkened; pistillode obsolete, resin scanty. 

bracteoles deltate, 1.8-2 mm long and wide; inner bracteoles oblate, 2.7- 

sepals orbicular, 6-7 mm long and wide; petals strongly cucuUate, 6-6.5 
mm long, 3.5-4 mm wide, staminodes 5, flat, 2.5-3-5 mm long, the fila- 
ment ca. 1 mm long, abruptly tapering at apex, the anther linear, ca. 1.5- 
2.5 mm long, apex broadly rounded, not well-differentiated from apex of 
filament, the thecae not fully differentiated and devoid of pollen, the con- 
nective darkened; pistil 3-5 mm long, 4-6 mm wide; carpels 5; styles 
absent, stigmas thinly coriaceous, suborbicular, peltate, 1.5-2 mm long 
and wide, thickening with age, flat in early ontogeny, convex at maturity; 
ovules numerous. Fruit yellowish-cream, suffused with purple along the 
carpel sutures, oblongoid, 2—3.5 cm long, 1.5—2.5 cm diam. 
Common name. — "Jubaga bakoko" (Mui language, Colombia) 
Distribution. — From lowlands of central Panama southward along the 
Pacific coast of Colombia (Choco Floristic Province), along and below the 
western slopes of the Andean Cordillera Occidental of Ecuador, eastward 
through the Amazon Basin of Colombia and Brazil, and south-southwest- 
ward through the Amazon Basin of Ecuador, Peru and Bolivia, 100-400 
(-1,690) m elevation. 

512 SiDA 16(3) 1995 

Ecology and conservation status. — Clusia amazonka is a riparian species, tol- 
erant of moderate amounts of disturbance but not capable of surviving 
long periods of indundation. It usually occurs high enough up in the canopy 
to permit some of the foliage to remain above periodic floodwaters. It oc- 
curs in varzea, igapo, tahuampa and other periodically inundated habitats. 
Unlike Clusia martiana, C. amazomca spends its entire life as a liana, and 
does not form large mats over the canopies of the trees. It is only very 
locally common, but is not considered threatened. 

Specimens examined. COLOMBIA. Amazonas: Rio Caqueta River Basin, 00°5()'S, 
7 1°50'W, 20 Nov 1991 (ster.),/ Dutvemmrden et al. 1259 (BRIT, COAH, COL, U); Mpio. 
Leticia, 03°47'S, 70°15'W, 100 m, 14 Apr 1991 (fr), /. Pm/y 15398 (BRIT, COL, FMB, 

HUA, JAUM, MO). VENEZUELA. Amazonas: 1 
near Culebra, 03°44'N, 65°44'W, 210 m, 15-16 Feb 1985 (stam. fl), R. Ltesner 17516 
(MO, VEN), (fr), R. Ltesner 17535 (BRIT, MO, VEN); Salto Yureba, Cano Yureba, Bajo 
Ventuari, 04°03'N, 66°01'W, 120-150 m, 4 Nov 1981 (scam, fl bud), F. Delasao & F. 
Gudnchez 10841 (MO, NY, US, VEN), 350 m, 15-16 Mar 1985 (pist. fl, fr), R. Liesner 
18755 (MO, VEN); Depto. Acurcs, near Remo Camp, 04°34'N, 67°18'W, 180 m, Jun 

1989 (fr), E. Foldats &J. Velaza, 9580 (MO, PORT, VEN); Depco. Casiquiare, 1 2 km SE of 
San Fernando de Atabapo, Sector "El Pozo," CVG Experiment Station, 03°50'N, 67°47'W, 
1 10 m, 10-16 Feb 1988 (]msc. fl), C. Aynuml et al. 6391 (MO, PORT, VEN); G. Aymard et 
al. 6404 (MO, PORT, VEN); Depto. Rio Negro, Cerro de la Neblina, vicinity of base 
camp along Rio Mawarinuma, afluent of Rio Baria, 00°50'N, 66°10'W, 1 10-130 m, 8-10 
Jan 1984 (stam. fl hud) J. Steyermark &J. Luteyn 129759 (F, MO, NY, US, VEN); Margins 
of Rio Gavilan, between Randal Gavilancito and Cerro Pelon, 05°37'N, 67°22'W, 80- 
120 m, 10-11 Feb 1992 (pist. fl), G. Romero et al. 2373 (AMES, MO, VEN). ECUADOR. 
Esmeraldas: Ecological Reserve Cococachi-Cayapas, Parroquia Luis Vargas Torres, Ri'o 
Santiago, Poce Salt Marsh, 00°49'N, 78M5'W, 250 m, 23-27 Oct 1993 (fr), M. Tirado et 
al. 539 (BRIT, QCNE, MO). Napo: Canton Orellana, Parque Nacional Yasuni, "Maxus" 
road and oil pipeline, km 20, 00°33'S, 76°30'W, 250 m, 11-15 Aug 1993 (pist. fl bud), 
M. Anlestia 306 (BRIT, MO, QCNE); Conoco oil well "Amo II," 00°52'S, 76°05'W, 230 
m, 29-30 Mar 1988 (fr), E Covello 152 (BRIT, MO, QCNE); Canton Tena, Jatun Sacha 
Biological Station, Rio Napo, 8 km E of Misahualli, 01°04'S, 77°36'W, 450 m, 20 Jan 

1990 (fr), C. Cerim 8386 (BRIT, MO, QCNE), 22-24 Feb 1988 (fr), W. Palaaos 2489 
(BRIT, MO, QCNE), 400 m, 17-28 May 1 989 (fr), W. Palaaos 4224 (BRIT, MO, QCNE), 
17-28 May 1989 (fr), W. Palacios 4307 (BRIT, MO, QCNE), 6 May 1990 (fr), W. Palacios 
etal. 4954 (BRIT, MO, QCNE), 13 May 1990 (fr), W. Palaaos & E. Freire 5105 (BRIT, 
MO, QCNE). 14 May 1990 (fr), W. Palaaos & E. Freire 5145 (BRIT, MO, QCNE), Perma- 
nent Inventory along Rio Chiguipino, 450 m, 22 May 1992 (fr), E. Guidtno &]. Zuleta 
1660 (BRIT, MO, QCNE). Pastaza: Canton Pastaza, "Masaramu" oil well, of UNOCAL, 
40 km NNW of Montalvo, 00°44'S, 76°52'W, 400 m, 1-16 May 1990 (fr), E. Guidino 
362 (BRIT, MO, QCNE); PetroCanada road. Via Auca, 115 km S of Coca, 5 km S of Rio 
Tiguino, 01°15'S, 76°55'W, 320 m, 26-31 Jan 1989 (fr), D. Neill & F Hurtado 8749 
(BRIT, MO, QCNE), 1-6 Mar 1989 (ster.), V.Zak4ll9 (BRIT, MO, QCNE). Sucumbios: 
Canton Gonzalo Pizarro, Campo Bermejo No. 6 Norte, 30 km NE of Lago Agrio, 00°l4'N, 
77°13'W, 1,050 m, 23 Mar 1990 (fr), C. Ceron 9227 (BRIT, MO, QCNE). PERU. Loreto: 
Prov. Maynas; Allpahuayo, Institute of Amazonian Investigations (IIAP) Experimental 

1991 (stam. fl), R. Vdsqmz & N. JaramUlo 14830 (AMAZ, BRIT, MO, USM), 27 May 
1991 (ster.), R- Vasquez & N.Jaramtllo 16618 (AMAZ, BRIT, MO, USM); Dtco. Fernando 
Lores, Caserfo Serafm, Quebrada Tamshiyacu, from Casen'o Serafin to Caserfo Constancia, 
125-130 m, 10 May 1991 (fr), C. Grdndez et al. 2606 (AMAZ, BRIT, MO, USM); Dtto. 
Iquitos, 1977 (stam. fl),/ Revilla 3312 (AMAZ, BRIT, F, MO, USM); Caseria Mjshana, 
Rio Nanay, right bank, 20 Dec 1976 (stam. fl),_/. Repilia 89 (AMAZ, F, MO, USM); Rio 
Yarapa, 04°20'S, 73°30'W, 122 m, 20 Nov 1989 (stam. fl bud), C. Grdndez &J. Tapullima 
1466 (AMAZ, MO, USM). BOLIVIA. Bern: Prov. Ballivian, Carinavi-San Borja Hwy, E 
side ofSerraniadel Pilon Lajas, 15°13'S, 67°03'W, 850 m, 2 Nov 1989 (fr), D. S?mth & Y. 
Garcia 13865 (LPB, MO). La Paz: Prov. Nor Yungas, near Yolosa, at entrance to road to 
Chairo, 2 km from Chairo, 2,430 m, 5 Sep 1987 (pist. fl), S. Beck 12969 (LPB, MO); Prov. 
Monllo, Valle de Zongo, along trail from end of road at the Cahua Power Plant, 1,660- 
1,690 m, 23 Nov 1980 (pist. fl), T. Croat 51413 (LPB, MO), 16°05'S, 68°03'W, 1,400- 
1,600 m, 22 Apr 1982 (fr),/ Sok?non 7491 (LPB, MO). BRAZIL. Amazonas: Alto Rio 
Solimoes, Mpio. de Sao Paulo de Oliven^a, plateau S of city, road to Bom Fim, 25 Nov 
1986 (stam. fl), C. Cid et al. 8532- A (BRIT, IAN, INPA, MG, NY). Para:Ilha do Marajo, 
RioMoc6es,2kmupriverfrom Anajas, ()0°57'S,49°56'W, 13 Nov 1987 (fr), 5. Beck et al. 
484 (BRIT, INPA, MG, NY), center of Iha Marajo, 27 Oct 1984 (stam. fl), G. SoM et al. 
4829 (BRIT, INPA, MG, NY). Rondonia : 28 km from Vilhena on road to Colorado, 
13°00'S, 60°00'W, 29 Oct 1979 (fr), B. Nelson et al. 330 (BRIT, IAN, INPA, MG, NY). 

Clusia amazonka is most closely related to C. araracuarae, but is readily 
distinguished by its smooth, obovate to elliptic leaf blades, muticous an- 
thers, and inconspicuous latex canals. This species is apparently sympatric 
with C. martiana, but occurs in riparian forests above and away from the 
inundation zone. Forest inventory data indicate that C. amazonka is much 
more infrequent than C. martiana, and does not form large mats as the 
latter species does. 

The type of C. oedematopoidea represents populations with smaller, less 
coriaceous leaves and smaller, more compact inflorescences than the Ama- 
zonian populations of C. amazonka. A collection from Antioquia, Colom- 
bia {R. Fonnegra et al. 4129) is identical to the type of C. oedematopoidea. 
Collections from Bolivia (i'. Beck 12969, T. Croat 31413, J. Solomon 7491) 
resemble C. martiana in quantitative features of the leaves, but the promi- 
nent secondary veins and submarginal collecting vein, and, absence of sig- 
nificant numbers of latex canals visible on the adaxial leaf surface, clearly 
indicate they belong to C. amazonica. 

3. Clusia martiana Engler, Fl. Bras. 12(l):4l 

■ngler, Bot. Jahrb. 58:Beibl 
lear Fortaleza, along Rio Jui 
5-destroyed, F Neg. 9205 !; 


epiphytic shrub, at times 

ippearing lianous t 

4 m tall 

branchlets te 

ete to somewhat tetragonal 

section, at 

times squarrc 

se, 2.5-3.5 mm diam., bark 

of apical portions ri 

fous, scaly 

checking crar 

sversely and exfoliate, rufou 

s and glabrous at ma 

urity, latex 

clear, moderate. Leaves decussate; blades 

ubcoriaceous, obova 

e to oblan- 

ceolate, (6.5- 

)10-14.5 (-19)cm long, (2. 


wide, apex 

narrowly acu 

e to abruptly acuminate, the acumen 0.3-1.5 cn- 

long, base 

acute to cune 

ite, not decurrent on the pet 

ole, smooth, glabrous above and 

below, sordid 

above, pallid below, latex ca 

als dense and conspicuous above 

and below, m 

drib slightly raised above, prominently raised bel 

3w, second- 

ary veins nur 

nerous, smooth, inconspicuc 

us above, slightly ra 

sed below 


■oUecting vein barely discern 

ible, the margin flat, 

entire, gla- 

brous; petioles canaliculate, 1-1 .5(-2) cm 

ong, glabrous. Inflor 

minal, pende 

nt, pyramidally paniculate. 

2-4 cm long, 3-5 

m wide at 

PiPOLY AND Grafi', Synopsis of Clusia 5 1 5 

maturity, the flowers 3 per cyme; peduncle 0.8-1.5 cm long, subtended by 
normal leaves; secondary bracts coriaceous, oblate, 2.3-2.6 mm long, 3.5- 
4 mm wide, apex widely rounded, carinate, the margin somewhat hyaline, 
very narrowly scarious, entire, glabrous. Staminate inflorescence with tet- 
ragonal pedicels, the lateral 1.8-2.2, the terminal 2.5-3-5 mm long, gla- 
brous; bracteoles 4, the outer 2 coriaceous, oblate, 2.3-2.6 mm long, 3.5- 

4 mm wide, apex widely rounded, carinate, the margin somewhat hyaline, 
very narrowly scarious, entire, glabrous, the inner 2 stiffly coriaceous, ob- 
late, 2.5-3 mm long, 3.3-4.5 mm wide, apex broadly rounded, somewhat 
carinate, the margins entire, somewhat translucent, not scarious, glabrous. 
Staminate flowers white to yellow; sepals 5, contorted, membranaceous, 
very widely ovate, 5-7 mm long, 4-6 mm wide, apex broadly rounded, 
somewhat cucuUate, densely lineate, the margin hyaline, scarious, glabrous; 
petals 5, contorted, carnose, oblong, 5.3-5.8 mm long, 1.8-2.2 mm wide, 
apex rounded, flat, obscurely lineate, the margin entire, opaque, not scari- 
otis, glabrous; androphore flat, pentagonal, the stamens numerous, free, 
2.1-3.3 mm long at maturity, the filaments short, flat, anther 1-1.3 mm 
long, apex muticous, base truncate, dehiscent by longitudinal slits, the 
connective flat, truncate, darkened; pistillode absent. Pistillate flowers white, 
as in staminate but sepals orbicular, 3-5-4 mm long and wide; petals 4.5- 

5 mm long, 2.8-3-3 mm wide, apex obtuse to rounded; staminodes 5, 3- 
3.5 mm long, the filaments wide, 1 mm long, 2 mm wide, abruptly con- 
stricted apically to anthers, the anthers linear, ca. 1 mm long, 0.5 mm 
wide, apex muticous, base not distinguishable, with longitudinal slits, de- 
void of pollen, the connective darkened; pistil 4 mm long, 5 mm diam.; 
carpels 5; styles absent, stigmas thinly coriaceous, oblongoid, peltate, 1.8- 
2.2 mm long, 1-1.2 mm wide, flat in early ontogeny, convex at maturity; 
ovules numerous. Fruit greenish white with red hue in intercarpelary areas, 
ovoid, 2-3-5 cm long, 1.5-2.5 cm wide. 

Distribution. — Amazon Basin of Colombia, Venezuela, Peru, Bolivia and 
Brazil, 100-450(-l,850) m elevation. 

Ecology and conservation status. — Clusia martiana is a hemiepiphyte, grow- 
ing along riverbanks, where it forms mats in and over the canopies of low 
trees, eventually killing them. This species can withstand extended peri- 
ods submerged. I have seen the fruits floating on rivers, but it is doubtful 
that dispersion by fish is significant, because of the large numbers of small 
birds I have seen feeding from open capsules, on the red ariUate seeds. It is 
commonly found on the margins of varzea, and "tahuampa" forests, which 
are on latentic soils, but is also rarely found in sandy loams ("varillal") and 
very rarely found in riparian forests with some pockets of white sand. Clusia 
martiana is one of the few weedy species oi Clusia. 

516 SiDA 16(3) 1995 

Specimens examined: COLOMBIA. Amazonas: Mpio. Leticia, Parque Nacionai Natu- 
ral Amacayacu, Matamata trail, ()3°47'S, 70°15'W, 110-120 m, 28 Oct 1991 (ster.),/ 
Pipoly 15610 (BRIT, COL, FMB, MO), near Quebrada Matamata, 100 m, 1 1 Mar 1991 
(fr), A. Ruclas et al. 1342 (BRIT, COL, FMB, MO), along Quebrada Bacaba, 100 m, 1 6 Apr 
1992 (fr), A. Ruclas & A. Prteto4286 (BRIT, COL, FMB, MO), along bank of Rio Amacayacu, 
100 m, 9 Apr 1991 (ster.),/ Pipoly et al. 15133 (BRIT, COL, FMB, MO); Vereda Puerto 
Narino, 03°45'S, 70°15'W, 100 m, 3 Aug 1989 (ster.), R. Vasquez et al. 12510 (AMAZ, 
BRIT, COL, FMB, MO, USM). Antioquia: Mpio. Amalfi, 8-27 km NE of Almafi, en 
route from Vetilla to Fraguas, near Salazar and Marengo, 06°00'N, 75°04'W, 1,150-1,450 
m, 7 Dec 1989 (pist. fl), R. Callejas et al. 9114 (BRIT, COL, HUA); Mpio. Campamento, 

75°20'W, 1,650-1,810 m, 7 Sep 1989 (pist. fl), R. Callejas et al. S266 (BRIT, COL, HUA).' 
VENEZUELA. Amazonas: Isla Sebastian, Ri'o Casiquiare above Chapezon, between Boca 
and Solano, 01°58'N,67°03'W, 120 m, 31 Jan 1980 (pist. fl), /?. Uesner&H. Clark 8941 
(MO, VEN); Depto. Atabapo, "Caiio Iguapo, Alto Orinoco, 15 km SE of La Esmeralda, 
03°00'N, 65°28'W, 150 m, 24 Feb 1990 (stam. fl), G. Aymard & L Delgado 8227 (BRIT, 
PORT, VEN); Depto. Atures, Rio Jenita floodplain, 7 km before Ocamo river mouth, 
02°46'N, 64°54'W, 170 m, Feb 1990 (stam. fl), A. Fernandez 7318 (BRIT, PORT, VEN); 
Rio Mayaca, 02°01'N, 65°07'W, 228 m, 6 Feb 1989 (stam. fl), A. Henderson et al. 987 
(BRIT, NY, VEN); Alto Orinoco, along riverside 2 hours above La Esmeralda, 26 Mar 
1953 (stam. fl), B. Magutre &J. Wurdack 34708 (MO, NY, VEN); Rio Padamo, 10 km 
above mouth, 27 Mar 1953 (stam. fl), B. Maguire &J. Wurdack 34718 (MO, NY, VEN); 
Depto. Casiquiare, near Capihuara, along Rio Casiquiare, 5 Feb 1991 (pist. fl, fr), M. 
Collela et al. 1829 (BRIT, NY, VEN); Isla Sebastian, along Rio Casiquiare between Boca 

(MO, VEN). PERU. Cuzco: Dtto. Camanti, 8 km W of Quincemil, right margin of Rio 

M. Trmand & H. Astete 646 (BRIT, CU2, MO, USM). Juni'n: Mazamari, 1,000 m, 13 Sep 
I960 (pist. fl, fr), F. Woytkowski 6019 (MO, US, USM). Loreto: Prov. Maynas, Buena Vista, 
Rio Tahuayo, 04°15'S, 73°10'W, 140 m, 24 Jan 1981 (stam. fl), R. Vasquez & N.J aramillo 
1229 (AMAZ, F, MO, USM); Explornapo Camp, Quebrada Sucusari, Rio Napo, 03°15'S, 
72°55'W, 130 m, 30 May 1991 (ster.), A. Gentry et al. 74277 (AMAZ, BRIT, MO, USM), 
Explornapo Camp, 03°20'S, 72°55'W, 140 m, 18 Apr 1991 (fr), R. Vasquez & N.J aramillo 
16141 (AMAZ, BRIT, MO, USM); Vicinity of Iquitos, 1977 (stam. fl),y. Revilla 3254 
(AMAZ, BRIT, F, MO, USM); Rio Mamon, 6 Sep 1972 (stam. fl), T. Croat 20028 (AMAZ, 
MO, USM); Santa Maria de Nanay, 10 km W of Caseria Mishana, Reserva Cocha Yarama' 
along Rio Nanay, 03°55'S, 73°35'W, 130 m, 15 Mar 1991 (ster.),/ Pipoly et al. 15031 
(AMAZ, BRIT, MO, USM), (fr),_/. Pipoly et al. 15036 (AMAZ, BRIT, CUZ, MO, USM); 
Rio Itaya, below San Juan de Muniches, ca. 40 min above Iquitos), 120 m, 19 Mar 1977 
(fr), A. Gentry et al. 18422 (AMAZ, F, MO, USM); Rio Nanay, 6 turns above Iquitos, 28 
Dec 1976 (pist. fl, fr),/ Revilla 2098 (AMAZ, F, MO, TEX, USM); Rfo Nanay between 
Iquitos and Puerto Almendras, 13 Jul 1976 (stam. fl), A. Gentry & J. Revilla 167 10 (AMAZ, 
F, MO, USM); Dtto. Iquitos, Rio Nanay, near Santa Clara, 19 Aug 1976 (stam. fl),/ 
Revilla 1144 (AMAZ, F, MO, USM), 13 Nov 1976 (stam. fl),/ Revilla 1899 (AMAZ, 
CUZ, F, HUT, MO, USM), 14 Nov 1976 (pist. fl bud),/ Revilla 1817 (AMAZ, F, MO, 
USM); Rio Nanay, near Santa Clara, ca. 10 km above Morona Cocha, 1 50 m, 12 Dec L976 
(stam. fl), C. Davidson &J. Revilla 5415 (AMAZ, F, MO, USM); Nauta, Quebrada Saragosa, 
04°29'S, 73°35'W, 200 m, 29 Mar 1987 (fr), R. Vasquez & N. Arevalo 9048 (AMAZ, MO, 

. Gentry &J. Revtlla 20814 (AMAZ, BRIT, CUZ, F, HUT, MO, TEX, USM); Along Rio 
lanco, above Tamshiyacu, 140 m, 17 Mar 1978 (fr), C. Diaz et al. 21^ (AMAZ, CUZ, F, 
lO, USM); Caserio Nina Rumi, along Rio Nanay, 23 Feb 1976 (fr),J. Revil/a 1 99 (AMAZ, 
[O, USM); San Antonio, Rio Pintuyacu, 03°40'S, 73°54'W, 160 m, 21 Apr 1986 (fr), R. 
chqiiez et al. 7487 (AMAZ, CUZ, F, MO, USM); Quebrada Yanayacu, tributary of Rio 

-a Paz: 1 


1, l4kmSWofIxiamas, 


I), A. Gentry & R. Foster 

km be 

low Yolosa, t 

hen 14 km W on road 

second ridgetop, 13°53'S, 68°15'W, 620-( 

70864 (F, LPB, MO); Prov. Nor Yungas, - 

along Ri'o Huarinilla, on abandonded road to Hacienda SandiUan, S side of river, 16°12'S, 

67°50'W, 1,200-1,300 m, 24 Jan 19S5 (fr) J. Solomon 9391 (LPB, MO); Prov. Munllo, 1/ 

2-3/4 by trail downriver from the Cahua Hydropower Plant, l6°05'S, 68°03'W, 1,400- 

1,600 m, 22 Apr 1982 (fr),J. Solomon 7491 (LPB, MO); 44 km below Lago Zongo Dam, 

vi'cmityofCahua hydroelectric plant, 16°03'S, 68°01'W, 1,200 m, 12-15Sep 1983(stam. 

fl),J. Solomon 10829 (LPB, MO). BRAZIL. Rorai. 

ma: SEMA Ecological Reserve, 

Ilha de 

Maraca, Parimiu, 03